taxonID	type	description	language	source
9A200C6C5D7A52C0BA6139507296112C.taxon	description	Figs 12 B, 15 A	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9A200C6C5D7A52C0BA6139507296112C.taxon	materials_examined	Type. Mexico. Chiapas: Mun. Cintalapa, Rancho El Cafetal, 950 m, 4 Mar 1996, M. A. Perez-Farrera 889 (holotype: CHIP! [acc. # 14306]; isotypes: F! [acc. # 2193633], HEM! [acc. # HEM 004830], MEXU! [MEXU 00827362], MO! [acc. # 04882667]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9A200C6C5D7A52C0BA6139507296112C.taxon	description	Description. Stem 20 - 60 cm long, 15 - 30 cm in diameter, epigeous, erect and decumbent. Cataphylls 2.0 - 5.0 x 1.5 - 3.0 cm wide at the base, persistent, triangular, reddish brown, densely brownish tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 2 - 30 (40), 36 - 125 cm long, ascending, brown at emergence with whitis gray trichomes, glabrous at maturity. Petiole 10 - 60 cm long, terete, linear, brown in mature leaves; with 11 - 40 thin prickles, 0.19 - 0.37 cm long. Rachis 25 - 87 cm long, terete, linear, brown to greenish brown in mature leaves, with prickles. Leaflets 25 - 68 pairs, opposite to subopposite, insertion in one plane, linear to lanceolate, generally longitudinally planar, not basally falcate, papyraceous, slightly involute, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetric at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 12 - 38 x 0.3 - 0.9 cm, 0.14 - 1.30 cm between leaflets; articulations 0.14 - 0.75 cm wide, brown. Pollen strobili 15 - 40 cm long, 3.5 - 5.1 cm in diameter, solitary, cylindrical, erect, yellowish green with brown trichomes at emergence, yellowish cream with blackish brown trichomes at maturity; peduncle 4 - 10 cm long, 1.5 - 2.1 cm in diameter, reddish brown to brown pubescent; microsporophylls 1.24 - 1.56 x 0.59 - 0.83 cm, elliptic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.50 - 0.81 cm long and linear with straight horns 0.14 - 0.30 cm long, 0.45 - 0.90 cm and an obtuse to acute angle between the horns. Ovulate strobili 17 - 27 cm long, 7.2 - 12.3 cm in diameter, solitary, cylindrical, erect, yellowish green with abundant blackish trichomes at emergence, glaucous green with reddish brown to blackish trichomes at maturity, acuminate apex; peduncle 4.0 - 10 cm long, 1.1 - 2.2 cm in diameter, erect, with scarce brownish tan trichomes; megasporophylls 20 - 80, 4 - 8 orthostichies with 5 - 10 sporophylls per orthostichy, 2.15 - 2.80 x 3.90 - 6.30 cm, with a prominent distal face, horns straight and thin and 0.32 - 0.51 cm long, 0.69 - 1.80 cm between horns with an obtuse angle between the horns. Seeds 2.5 - 3.0 cm long, 2.3 - 2.9 cm in diameter, spherical, sarcotesta whitish pink when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9A200C6C5D7A52C0BA6139507296112C.taxon	distribution	Distribution and habitat. Ceratozamia alvarezii is endemic to Mexico and only known from Cintalapa and Jiquipilas municipalities in Chiapas State, at the transition zone between pine and oak forest and oak forest; plants occur on karstic rocks between 900 and 1,450 m elevation (Fig. 14 A).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9A200C6C5D7A52C0BA6139507296112C.taxon	etymology	Etymology. The specific epithet honors Miguel Alvarez del Toro in recognition of its outstanding contributions for conservation in Chiapas and the establishment of its first reserves (Perez-Farrera et al. 1999).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9A200C6C5D7A52C0BA6139507296112C.taxon	discussion	Discussion. The morphology of Ceratozamia alvarezii is not very different from C. mirandae. At population level, this species differs from C. mirandae by having generally shorter leaves, smaller ovulate strobilus (up to 27 cm long and 5 to 10 sporophylls per orthostichy), and longer seeds.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9A200C6C5D7A52C0BA6139507296112C.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Mun. Cintalapa, 1,100 m, 10 Mar 1993, A. P. Vovides 1234 (XAL); 920 m, 21 Jun 2018, F. Nicolalde-Morejon et al. 2791 - 2799 (CIB); 980 m, 22 Jun 2018, F. Nicolalde-Morejon et al. 2830 (CIB); 1,107 m, 18 Jun 2019, F. Nicolalde-Morejon & L. Martinez-Dominguez 3177 - 3183 (CIB); 1,350 m, 18 Jun 2019, F. Nicolalde-Morejon & L. Martinez-Dominguez 3186 - 3196 (CIB); 1,450 m, 11 Oct 1994, J. Castillo et al. 445 (CHIP); 920 m, 21 Jun 2018, L. Martinez-Dominguez et al. 1359 - 1369 (CIB), 1370 (CIB, MEXU); 980 m, 22 Jun 2018, L. Martinez-Dominguez et al. 1402 (CIB, MEXU); 925 m, 18 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1770 (CIB); 1,107 m, 18 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1771,1772 (CIB, MEXU), 1773 (CIB), 1774 (CIB, MEXU), 1775 - 1777 (CIB); 1,350 m, 18 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1784 - 1794 (CIB); 900 m, 10 Jul 1994, M. A. Perez-Farrera 71 (CIB, CHIP, MEXU); 950 m, 5 Sep 1995, M. A. Perez-Farrera 776 (CIB, XAL); 1,000 m, 27 Mar 2000, O. Farrera S. 2068 (CHIP). Mun. Jiquipilas, 1,200 m, 6 Jun 2002, A. Reyes-Garcia 5017 (MEXU); 1,380 m, 17 Feb 2000, E. Palacios E. 2469 (CHIP); 1,170 m, 7 Jul 1994, M. A. Perez-Farrera 68 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5E0430B3E8B75713A3465D69A948237E.taxon	description	Figs 1 C, 15 B	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5E0430B3E8B75713A3465D69A948237E.taxon	materials_examined	Type. Mexico. Oaxaca: Mun. San Pedro Teutila, Sierra Norte, 30 May 2021, M. A. Perez-Farrera & P. Diaz-Jimenez 4014 (holotype: HEM [n. v.]; isotype: XAL [n. v.]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5E0430B3E8B75713A3465D69A948237E.taxon	description	Description. Stem 30 - 250 cm long, 10 - 15 cm in diameter, epigeous, erect or sometimes decumbent. Cataphylls 3.6 - 6.5 x 2 - 3.3 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 7 - 30 (50), 117 - 240 cm long, ascending, reddish brown at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 45 - 85 cm long, terete, linear, reddish brown or greenish brown at emergence, dark green in mature leaves; with 16 - 47 thin prickles, 0.30 - 0.66 cm long. Rachis 70 - 160 cm long, terete, linear, dark green in mature leaves, with prickles in lower third. Leaflets 12 - 38 pairs, opposite to subopposite, insertion in one plane, oblong, generally longitudinally planar, not basally falcate, papyraceous, flat, green with adaxial and abaxial sides glabrous, distal end with entire margins, abruptly acuminate and symmetrical at the apex, attenuate at base, veins conspicuous and light-green; median leaflets 21 - 42.5 x 3.3 - 4.8 cm, 3.2 - 6.0 cm between leaflets; articulations 0.9 - 1.5 cm wide, green. Pollen strobili 20 - 35 cm long, 3.1 - 4.6 cm in diameter, solitary, cylindrical, erect, greenish with reddish trichomes at emergence, greenish yellow with reddish brown trichomes at maturity; peduncle 7.0 - 15 cm long, 1.5 - 2.1 cm in diameter, pubescent, reddish brown; microsporophylls 1.50 - 2.2 x 0.98 - 1.30 cm, obconic with a non-recurved distal face and a lobate fertile portion, infertile portion rounded and 0.47 - 0.84 cm long with horns straight and 0.16 - 0.38 cm long, 0.50 - 0.80 cm between horns with an acute angle between the horns. Ovulate strobili 11 - 36 cm long, 9.4 - 12.5 cm in diameter, solitary, cylindrical, erect, greenish yellow with an abundant reddish trichomes at emergence, green with brown trichomes at maturity, with an acute apex; peduncle 5.0 - 19.5 cm long, 1.3 - 2.3 cm in diameter, erect, light brown pubescent; megasporophylls 20 - 200, 5 - 11 orthostichies with 4 - 19 sporophylls per orthostichy, 2.25 - 2.72 x 3.6 - 4.3 cm, with a truncate distal face, horns curved and 0.54 - 0.90 cm long, 1.05 - 1.77 cm between horns with an acute angle between the horns. Seeds 2.4 - 3.0 cm long, 1.4 - 1.7 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5E0430B3E8B75713A3465D69A948237E.taxon	distribution	Distribution and habitat. Ceratozamia aurantiaca occurs in highlands south of Rio Santo Domingo and north of Sierra Norte in Oaxaca State, Mexico (Fig. 14 B), where it occurs in the evergreen tropical forest on karstic rocks at 458 - 800 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5E0430B3E8B75713A3465D69A948237E.taxon	etymology	Etymology. The specific epithet alludes to the leaf color at emergence. This is derived from Latin word aurantiacus for the orange color of emerging leaves.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5E0430B3E8B75713A3465D69A948237E.taxon	discussion	Discussion. Ceratozamia aurantiaca is most similar to C. whitelockiana, however, differs from it in its oblong leaflets abruptly acuminate with a symmetric apex, its petiole with abundant (16 - 24) and long prickles (0.32 - 0.55 cm), and its rachis that has prickles in the proximal third of the leaf. In addition, the ovulate strobili have an acute apex, and the pollen strobili have obconic microsporophylls with a rounded infertile portion. The leaves are reddish brown at emergence, whereas in C. whitelockiana they are green.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5E0430B3E8B75713A3465D69A948237E.taxon	materials_examined	Specimens examined. Mexico. Oaxaca: Mun. Chiquihuitlan de Benito Juarez, 861 m, 26 Nov 2004, C. A. Cruz-Espinosa & G. Juarez-Garcia 1945 (MEXU). Mun. San Felipe Jalapa de Diaz, 500 m, 28 Jun 2008, J. A. Perez de la Rosa et al. 1966 (IBUG). Mun. San Pedro Teutila, 458 m, 23 Sep 2020, F. Nicolalde-Morejon et al. 3323 - 3333 (CIB); 500 m, 23 Sep 2020, F. Nicolalde-Morejon et al. 3334 - 3337 (CIB); 615 m, 28 Sep 2020, F. Nicolalde-Morejon et al. 3415 - 3419 (CIB); 708 m, 10 Jun 2004, G. Juarez-Garcia 425 (MEXU); 458 m, 23 Sep 2020, L. Martinez-Dominguez et al. 1946 - 1950 (CIB), 1951 (CIB, MEXU), 1952 - 1957 (CIB); 500 m, 23 Sep 2020, L. Martinez-Dominguez et al. 1958 - 1960 (CIB, MEXU), 1961 (CIB); 615 m, 28 Sep 2020, L. Martinez-Dominguez et al. 2050, 2052 (CIB, MEXU), 2051, 2053 (CIB); 539 m, 6 Apr 2021, L. Martinez-Dominguez et al. 2141, 2142, 2144 (CIB), 2143 (CIB, MEXU, NY). Mun. Santa Maria Tlalixtac, 675 m, 25 Nov 2004, G. Juarez-Garcia & C. A. Cruz-Espinosa 868 (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
960BDC31034E58088BBCA73C600FFCAC.taxon	description	Fig. 15 C	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
960BDC31034E58088BBCA73C600FFCAC.taxon	materials_examined	Type. Mexico. Tabasco: Mun. Teapa, hill Madrigal, Aug 2003, A. P. Vovides 1458 (holotype: XAL [XAL 0148355]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
960BDC31034E58088BBCA73C600FFCAC.taxon	description	Description. Stem 13 - 38 cm long, 9 - 15 cm in diameter, epigeous, semi-hypogeous, erect. Cataphylls 2.6 - 5.7 x 1.3 - 4.5 cm wide at the base, persistent, triangular, reddish brown, densely brownish tomentose at emergence, glabrous at apex when mature, apex acuminate. Leaves 2 - 12 (17), 49 - 210 cm long, descending, light green, glaucous at emergence with whitish gray trichomes, glabrous at maturity. Petiole 35 - 111.5 cm long, terete, linear, pink at emergence, yellowish green when mature; unarmed to armed with 3 - 27 thin prickles, 0.02 - 0.13 cm long. Rachis 30 - 105 cm long, terete, linear, pink at emergence, yellowish green at mature leaves, generally unarmed. Leaflets 6 - 15 pairs, opposite to subopposite, insertion in one plane, oblong to oblanceolate, longitudinally curved abaxially to planar, generally basally falcate, coriaceous, flat, green with adaxial and abaxial sides glaucous or glabrous, distal end with entire margins, acuminate to acute, and asymmetrical (rarely symmetrical in apical leaflets) at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 16 - 30 x 4.5 - 8.8 cm, 5.5 - 11.1 cm between the leaflets; articulations 0.5 - 1.5 cm wide, green and yellowish. Pollen strobili solitary, 10.8 - 20 cm long, 2.8 - 4.0 cm in diameter, cylindrical, erect, green with blackish trichomes at emergence, yellow-cream with blackish trichomes at maturity; peduncle 5 - 10 cm long, 1.5 - 2.0 cm in diameter, tomentose, light brown; microsporophylls 0.9 - 1.37 x 0.7 - 0.9 cm, discoid with a non-recurved distal face and a deeply lobate fertile portion, infertile portion 0.35 - 0.39 cm long and linear with straight horns and 0.30 - 0.45 cm long, 0.70 - 0.80 cm and an acute angle between the horns. Ovulate strobili 15 - 23 cm long, 7.5 - 8.5 cm in diameter, solitary, cylindrical, erect, green with reddish brown trichomes at emergence, green with brown to blackish trichomes at maturity, acute apex; peduncle 5 - 12 cm long, 1.6 - 2 cm in diameter, pendulous and erect, tomentose, light brown; megasporophylls 43 - 56, 6 - 8 orthostichies with 5 - 7 sporophylls per orthostichy, 1.7 - 2.0 x 3.5 - 4.0 cm, with a prominent distal face, horns straight and 0.70 - 0.87 cm long, 0.92 - 1.50 cm between horns with a right angle between the horns. Seeds 1.5 - 2.4 cm long, 1.2 - 2.0 cm in diameter, ovate, sarcotesta whitish pink at emergence, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
960BDC31034E58088BBCA73C600FFCAC.taxon	distribution	Distribution and habitat. Ceratozamia becerrae is endemic to the mountains of Sierra Madrigal in southern Tabasco and Chiapas States, Mexico (Fig. 14 C), where it occurs on karstic outcrops in evergreen tropical forest and oak forest from 100 - 800 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
960BDC31034E58088BBCA73C600FFCAC.taxon	etymology	Etymology. The specific epithet was established in honor of Professor Marco E. Becerra for his relevant contributions in ethnology, archaeology and floristic research in Tabasco (Vovides et al. 2004).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
960BDC31034E58088BBCA73C600FFCAC.taxon	discussion	Discussion. Ceratozamia becerrae has oblong and coriaceous leaflets and leaves that have a few thin and short prickles. This species belongs to a cryptic taxonomic group with C. zoquorum and C. santillanii (c. f. Martinez-Dominguez et al. 2017 c; Vovides et al. 2020) and is morphologically similar in both vegetative and reproductive characters to C. zoquorum. This species differs from C. santillanii by its peduncle of ovulate strobili more than 3 cm long, but lacks morphological diagnoses with C. zoquorum.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
960BDC31034E58088BBCA73C600FFCAC.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Mun. Amata ́ n, 300 m, J. M. La ́ zaro Z. 376 (CHIP). Tabasco: Mun. Tacotalpa, 260 m, 12 Apr 2014, F. Nicolalde-Morejo ́ n et al. 1999, 2000 (CIB); 260 m, 12 Apr 2014, L. Marti ́ nez-Domi ́ nguez et al. 108 - 127 (CIB); 19 Jan 2001, S. Avendan ̃ o R. 5214 b (XAL). Mun. Teapa, 100 m, 29 Jan 1985, B. M. Schutzman 645, 648 - 650 (XAL); 204 m, 22 Feb 2014, F. Nicolalde-Morejon et al. 1968, 1969 (CIB); 800 m, 11 Jun 1989, J. A. Alejandre Rosas 494 (CIB); 204 m, 22 Feb 2014, L. Marti ́ nez-Domi ́ nguez et al. 35 - 40 (CIB); 400 m, 16 Apr 1996, M. A. Pe ́ rez-Farrera 901 (CHIP, HEM, MEXU); 800 m, 7 Apr 1914, M. E. Becerra s / n (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
8D2FCB5EFA525DE3AD0DAEBDF3537D6E.taxon	description	Figs 3 B, 12 C, 14 D, 15 D	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
8D2FCB5EFA525DE3AD0DAEBDF3537D6E.taxon	materials_examined	Type. Mexico. Veracruz: Mun. Alto Lucero de Gutierrez Barrios, Apr 2005, S. Avendan ̃ o R. 5699 (neotype, designated by Vovides et al. 2012, pg. 38: XAL! [XAL 0132508]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
8D2FCB5EFA525DE3AD0DAEBDF3537D6E.taxon	description	Description. Stem 20 - 70 cm long, 15 - 40 cm in diameter, epigeous, erect. Cataphylls 2 - 5 x 1.5 - 4 cm wide at the base, persistent, triangular, reddish brown, densely brownish tomentose at emergence, glabrous at maturity with an acuminate apex. Leaves 6 - 36, 58 - 173.5 cm long, descending, yellowish green at emergence with brown trichomes, glabrous at maturity. Petiole 20 - 56 cm long, terete, linear, green in mature leaves; with 40 - 90 robust prickles, 0.24 - 50 cm long. Rachis 35 - 125.5 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 13 - 38 pairs, opposite to subopposite, insertion keeled, sometimes imbricate, lanceolate, abaxially curved, basally falcate, coriaceous, flat, light green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical to asymmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 15.5 - 41 x 2 - 4.1 cm, 0.5 - 3.2 cm between leaflets; articulations 0.6 - 1.7 cm wide, yellow. Pollen strobili 18 - 31 cm long, 4 - 7 cm in diameter, solitary, cylindrical, erect, greenish yellow at emergence, greenish yellow with brown to blackish trichomes at maturity; peduncle 4 - 10.3 cm long, 1.7 - 2.2 cm in diameter, reddish brown to light-brown pubescent; microsporophylls 1.55 - 2.3 x 0.80 - 1.5 cm, obconic with a non-recurved distal and a lobate fertile portion, infertile portion 0.50 - 0.70 cm long and rounded with straight horns 0.24 - 0.40 cm long, 0.50 - 1.02 cm and an acute angle between the horns. Ovulate strobili 25 - 33 cm long, 9.8 - 12.5 cm in diameter, solitary, cylindrical, erect, green with blackish trichomes at emergence, greenish yellow with brown to blackish trichomes at maturity and with an acuminate and apiculate apex; peduncle 6 - 14 cm long, 2.0 - 2.4 cm in diameter, pendulous and erect, brown to reddish brown pubescent; megasporophylls 80 - 224, 8 - 16 orthostichies with 10 - 15 sporophylls per orthostichy, 1.5 - 2.8 x 2.5 - 3.2 cm, with a prominent distal face, horns straight and robust and 0.75 - 0.95 cm long, 1.15 - 1.70 cm between horns with an acute angle between the horns. Seeds 2.0 - 3.0 cm long, 1.3 - 2.0 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
8D2FCB5EFA525DE3AD0DAEBDF3537D6E.taxon	distribution	Distribution and habitat. Ceratozamia brevifrons is known only from Sierra de Chiconquiaco in Veracruz State, Mexico (Fig. 14 D) where it occurs in the transition zone between cloud forest and oak forest at 450 to 1,370 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
8D2FCB5EFA525DE3AD0DAEBDF3537D6E.taxon	etymology	Etymology. The epithet is derived from its relatively short leaves.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
8D2FCB5EFA525DE3AD0DAEBDF3537D6E.taxon	discussion	Discussion. Ceratozamia brevifrons is easily distinguished from its congeners by having adaxially keeled and coriaceous leaflets, petioles armed with abundant short and robust prickles and greenish yellow ovulate strobili with brown to blackish brown trichomes at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
8D2FCB5EFA525DE3AD0DAEBDF3537D6E.taxon	materials_examined	Specimens examined. Mexico. Veracruz: Mun. Alto Lucero de Gutierrez Barrios, 24 Aug 1976, A. P. Vovides 119 (XAL); 8 Jan 2009, D. Jimeno-Sevilla 694 (XAL); 1,052 m, 12 Jan 2013, F. Nicolalde-Morejon et al. 1711 - 1731 (CIB); 1,052 m, 22 Aug 2014, F. Nicolalde-Morejon & L. Martinez-Dominguez 2027 - 2046 (CIB); 1,250 m, 6 Apr 1981, G. Castillo-Campos 1297 (XAL); 700 m, 3 Dec 1974, J. Rees 1636 (MO, XAL), 1641, 1642, 850 m, 21 Sep 1976, 1675 (XAL); 1,052 m, 21 Jun 2014, L. Martinez-Dominguez & F. Nicolalde-Morejon 130 - 133 (CIB); 842 m, 6 Feb 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 216 - 226 (CIB); 1,052 m, 22 Mar 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 298 - 309 (CIB); 450 m, 14 Jul 1995, M. Vazquez-Torres 4790 (CIB); 24 Jun 2010, M. Vazquez-Torres et al. 9186 (CIB); 850 m, 10 Jan 2001, T. W. Walters 2001 - 02 - A, B (XAL). Mun. Chiconquiaco, 1,268 m, 10 April. 2015, F. Nicolalde-Morejon & L. Martinez-Dominguez 2237 - 2241 (CIB); 1,340 m, 2 May 2019, F. Nicolalde-Morejon et al. 3138 - 3147 (CIB); 1,268 m, 10 Apr 2015, L. Martinez-Domiguez & F. Nicolalde-Morejon 556 - 560 (CIB); 1,340 m, 2 May 2019, L. Martinez-Domiguez et al. 1729 - 1738 (CIB). Mun. Colipa, Marts 1841 (MO). Mun. Juchique de Ferrer, 850 m, 30 Aug 1981, A. P Vovides 682 (XAL); 1,250 m, 6 May 1981, G. Castillo-Campos 1710, 1763, 1768 (XAL); 1,300 m, 7 May 1981, G. Castillo-Campos 1815, 1824, 1981 (XAL); 1,370 m, 24 Jul 2008, M. Vazquez-Torres 8633 (CIB). Mun. Vega de Alatorre, 650 m, 21 Jul 1981, B. Guerrero & J. I. Calzada 1826 (XAL); 550 m, 21 Jul 1981, G. Castillo-Campos 2033 (XAL).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3884E28F65AC57199CD99EF5F189E17E.taxon	description	Description. Stem 10 - 250 cm long, 8 - 40 cm in diameter, epigeous or semi-hypogeous, erect or decumbent. Cataphylls persistent, triangular to narrowly triangular, reddish brown, tomentose to densely tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves stipulate, ascending to descending, light green or reddish brown at emergence with whitish gray or brown trichomes, generally glabrous at maturity; stipulate 2 - 6 cm long, linear, tomentose at maturity. Petiole straight or twisted, sometimes brown in mature leaves, without prickles or heavily to lightly armed with prickles; prickles can be bifurcate. Rachis straight or twisted, without prickles or armed with prickles up to half the length of the leaves. Leaflets articulate, sessile, membranaceous to coriaceous, linear to obovate, opposite to subopposite or clustered, not imbricate, generally acuminate at apex, attenuate at base, margins entire; articulations green to brown. Pollen strobili 1 - 2, with sterile tip, erect, cylindrical, green to cream with blackish to reddish brown trichomes at maturity; pollen sporangiophores deltoid to cuneate, basally stalked, distal face bicornate, fertile abaxial surface with 24 - 280 sporangia in clusters of (2) 3 (4 - 5), dehiscent by longitudinal slit; peduncle pubescent to tomentose. Ovulate strobili usually solitary, globose to cylindrical; green with pale pink to blackish trichomes at maturity, acute to apiculate apex; ovulate sporangiophores peltate with a narrow basal stalk and transversely hexagonal tips, bicornate at distal end; peduncle pubescent to tomentose, erect to pendulous. Seeds (ovules) 2 per megasporophyll projecting inward toward the strobilus axis, spherical, sarcotesta pink to yellowish when immature, light brown at maturity, sclerotesta smooth with several furrows longitudinal from micropylar end.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3884E28F65AC57199CD99EF5F189E17E.taxon	distribution	Distribution and habitats. The 36 species of Ceratozamia are only found from Mexico to Central America, usually in montane habitats on limestone soils at elevations from 19 to 2,000 m. Most of the species are narrowly endemic, and all are on CITES Appendix I.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2626037019595B50B5DFF05114953E32.taxon	description	Figs 3 C, 15 E	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2626037019595B50B5DFF05114953E32.taxon	materials_examined	Type. Mexico. San Luis Potosi: Mun. Xilitla, 1,044 m, 20 Mar 2016, L. Martinez-Dominguez et al. 933 ♀ (holotype: CIB! [acc. # 17766 UV]; isotypes: MEXU! [MEXU 1492226, MEXU 1492227, MEXU 14922278], NY!).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2626037019595B50B5DFF05114953E32.taxon	description	Description. Stem 20 - 60 cm long, 15 - 30 cm in diameter, epigeous, erect and decumbent. Cataphylls 2.5 - 5.5 x 1.5 - 3.5 cm wide at the base, persistent, narrowly triangular, reddish brown, densely brownish tomentose at emergence, glabrous at maturity, apex acuminate. Leaves (5) 10 - 40, 100 - 207 cm long, descending, reddish brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole 30 - 69 cm long, terete, linear, blackish brown in mature leaves; with 9 - 30 robust prickles, 0.03 - 0.30 cm long. Rachis 65 - 144 cm long, terete, linear, reddish brown in mature leaves, with prickles. Leaflets 20 - 42 pairs, opposite to subopposite, insertion in one plane, oblong, longitudinally curved abaxially to planar, not basally falcate, coriaceous, flat, dark green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and asymmetric at the apex, attenuate at base, with conspicuous and reddish brown veins; median leaflets 20 - 37 x 2.3 - 4 cm, 1.6 - 3.5 cm between leaflets; articulations 0.5 - 1.3 cm wide, generally reddish brown. Pollen strobili 20 - 31 cm long, 4.5 - 6 cm in diameter, generally solitary (1 - 2), cylindrical, erect, greenish brown at emergence with reddish trichomes becoming greenish with reddish brown trichomes at maturity; peduncle 5.2 - 8.1 cm long, 1.6 - 2.3 cm in diameter, reddish brown pubescent; microsporophylls 1.6 - 2.3 x 0.9 - 1.5 cm, obconic with a non-recurved distal face and a deeply lobate fertile portion, infertile portion 0.44 - 0.55 cm long and linear with curved horns 0.25 - 0.40 cm long, 0.40 - 0.80 cm and an acute angle between the horns. Ovulate strobili 25.5 - 30 cm long, 7.5 - 10.5 cm in diameter, cylindrical, erect, greyish green with reddish brown trichomes at emergence and becoming light grayish brown with reddish brown trichomes at maturity with an acuminate apex; peduncle 4.5 - 11.5 cm long, 1.3 - 2.5 cm in diameter, erect, tomentose, brown to reddish brown; megasporophylls 49 - 180, 7 - 12 orthostichies with 7 - 15 sporophylls per orthostichy, 2.0 - 3.0 x 2.3 - 4.0 cm, with a prominent distal face, horns straight and 0.40 - 0.80 cm long, 2.0 - 2.70 cm between horns with an acute angle between the horns. Seeds 2.2 - 3.5 (4) cm long, 0.6 - 1.7 cm in diameter, ovate, sarcotesta whitish red to pink when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2626037019595B50B5DFF05114953E32.taxon	distribution	Distribution and habitat. Ceratozamia chamberlainii is distributed in the northern montane region of Carso Huasteco in San Luis Potosi ́, Quere ́ taro and Hidalgo States, Mexico (Fig. 16 A), where it occurs in cloud forest and pine-oak forest on rocky outcrops between 900 - 1,200 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2626037019595B50B5DFF05114953E32.taxon	etymology	Etymology. The specific epithet is in honor of Charles Joseph Chamberlain in recognition of his remarkable contributions to knowledge of the biology of the cycads and his fieldwork on Mexican Cycads (Martinez-Dominguez et al. 2017 a).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2626037019595B50B5DFF05114953E32.taxon	discussion	Discussion. Ceratozamia chamberlainii is close geographically and morphologically to C. fuscoviridis and C. latifolia; however, it differs by its petioles and rachis bearing abundant and short prickles, and its oblong and coriaceous leaflets with conspicuous, reddish brown veins. In addition, the leaves are reddish brown at emergence and at maturity have this color in the leaflet articulations and at the base of leaflets in contrast to the yellowish to greenish articulations in C. fuscoviridis and C. latifolia.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2626037019595B50B5DFF05114953E32.taxon	materials_examined	Specimens examined. Mexico. Hidalgo: Mun. Chapulhuacan, 1,500 m, 21 Feb 1998, Alcantara-Ayala 3650 (FCME); 8 Jan 2009, A. Vite-Reyes et al. 23 (XAL); 1,157 m, 30 Mar 2015, F. Nicolalde-Morejon et al. 2200 - 2203 (CIB); 20 Sep 1964, L. Gonzalez-Quintero 4634 (ENCB); 1,157 m, 30 Mar 2015, L. Martinez-Dominguez et al. 429 - 454 (CIB). Mun. La Mision, 1,120 a 1,400 m, 5 Oct 2007, A. Castro-Castro et al. 1017 (IBUG, XAL); 7 Jan 2009, A. Vite-Reyes et al. 20 (XAL). Queretaro: Mun. Landa de Matamoros, 1,050 m, 28 May 1999, A. P. Vovides 1288, 1289, 1290, 2000 (XAL), 1291 (XAL, MEXU); 2 Apr 1991, E. Carranza 6333 (IEB); 940 m, 2 Apr 1991, 3119 (MEXU); 1,335 m, 30 Mar 2015, F. Nicolalde-Morejon et al. 2192 - 2197 (CIB); 1,145 m, 30 Mar 2015, F. Nicolalde-Morejon et al. 2198, 2199 (CIB); 1,335 m, 30 Mar 2015, L. Martinez-Dominguez et al. 379 (CIB, MEXU), 380 - 407 (CIB); 1,145 m, 30 Mar 2015, L. Martinez-Dominguez et al. 408 - 428 (CIB); 1,070 m, 9 May 1989, R. Hiram 647 (XAL). San Luis Potosi: Mun. Aquismon, 600 m, 10 Feb 1969, H. Puig 3979 (ENCB, P); 27 May 1979, J. A. Alcorn 3093 (MEXU); 1,125 m, 9 Jun 2015, T. Diego-Vargas & M. Bonta 35 (XAL). Mun. Tamazunchale, Jul 1952, A. Wilson 273, 274 (US); Jul 1937, C. L. Lundell & A. A. Lundell 7235 (CIB, IEB); 600 - 900 m, 29 Jun 1959, J. Rzedowski 11087 (ENCB). Mun. Xilitla, 1,044 m, 20 Mar 2016, F. Nicolalde-Morejon et al. 2407 - 2420 (CIB); 1,044 m, 20 Mar 2016, L. Martinez-Dominguez et al. 924 - 932, 934 - 937 (CIB), 938 (CIB, MEXU), 939 (CIB); 1,948 m, 12 Jan 2001, T. W. Walters, TW- 2001 - 04 - A, B (XAL), TW-. 2001 - 04 - C (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
F5DCCDFED99051DAB219ACF618415FE8.taxon	description	Fig. 15 F	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
F5DCCDFED99051DAB219ACF618415FE8.taxon	materials_examined	Type. Mexico. Oaxaca: Chimalapa, 21 Jan 2002, M. A. Perez-Farrera 2622 ♀ (holotype: HEM [n. v.]; isotypes: XAL [XAL 0146074], MEXU [n. v.], MO [n. v.]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
F5DCCDFED99051DAB219ACF618415FE8.taxon	description	Description. Stem 20 - 60 cm long, 15 - 30 cm in diameter, epigeous, erect and decumbent. Cataphylls 4 - 7.5 x 2.5 - 4.3 cm wide at the base, persistent, triangular, reddish brown, scarce brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 7 - 24, 66 - 250 cm long, ascending, brown at emergence, glabrous at maturity. Petiole 40 - 70 cm long, terete, linear, green in mature leaves; with 20 - 30 thin prickles, 0.05 - 0.30 cm long. Rachis 60 - 150 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 36 - 70 pairs, opposite to subopposite, insertion in one plane, lanceolate, longitudinally curved abaxially to planar, not basally falcate, papyraceous, flat, light green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 27 - 46 x 1.2 - 1.8 cm, 0.8 - 1.3 cm between leaflets; articulations 0.5 - 0.9 cm wide, brown to yellowish brown. Pollen strobili 25 - 40 cm long, 3 - 5 cm in diameter, solitary, cylindrical, erect, greenish yellow at emergence with few brown trichomes, greenish to cream at maturity; peduncle 5 - 11 cm long, 1.5 - 3 cm in diameter, reddish brown pubescent; microsporophylls 1.3 - 2 x 0.6 - 1 cm, elliptic with a non-recurved distal face and a lobate to slightly lobate fertile portion, infertile portion 0.7 - 1.1 cm long and linear with curved horns 0.30 - 0.43 cm long, 0.8 - 1.1 cm and an acute angle between the horns. Ovulate strobili 35 - 40 cm long, 7.3 - 10.6 cm in diameter, solitary, cylindrical, erect, green with blackish trichomes at emergence, green with blackish trichomes at maturity, acuminate apex; peduncle 7 - 11 cm long, 1.7 - 2 cm in diameter, erect, brown to reddish brown pubescent; megasporophylls 99 - 192, 11 - 12 orthostichies with 9 - 16 sporophylls per orthostichy, 4 - 5.9 x 1.8 - 2.5 cm, with a prominent distal face, horns straight and thin and 0.78 - 0.98 cm long, 0.98 - 2.0 cm between horns with an acute angle between the horns. Seeds 2.0 - 3.0 cm long, 1.4 - 1.9 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
F5DCCDFED99051DAB219ACF618415FE8.taxon	distribution	Distribution and habitat. Ceratozamia chimalapensis is endemic to Oaxaca State (Mexico) in the Sierra Atravesada mountain range at 290 - 1,000 m (Fig. 16 B) in oak forest on clay soils.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
F5DCCDFED99051DAB219ACF618415FE8.taxon	etymology	Etymology. The specific epithet is in honor of the Chimalapa region renowned for its biological richness, particularly its floristic diversity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
F5DCCDFED99051DAB219ACF618415FE8.taxon	discussion	Discussion. Ceratozamia chimalapensis is similar to C. mirandae and C. alvarezii in leaf morphology; however, there are differences in reproductive structures; the ovulate strobilus of C. chimalapensis is longer than in these species. The ovulate strobili have between 11 - 12 orthostichies with 9 - 16 sporophylls per orthostichy, and more than 90 seeds per strobilus. The seeds are ovate, whereas in the other two species, they are spherical.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
F5DCCDFED99051DAB219ACF618415FE8.taxon	materials_examined	Specimens examined. Mexico. Oaxaca: Mun. Santa Maria Chimalapa, 290 m, 11 Jun 1995, R. Garcia S. 319 (SERO, XAL). Mun. Santiago Niltepec, 1000 m, 3 Apr 1946, E. Hernandez Xolocotzi & A. J. Sharp 1277 (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3E10BF94F81559809A078DF5A3A280B1.taxon	description	Fig. 15 G	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3E10BF94F81559809A078DF5A3A280B1.taxon	materials_examined	Type. Mexico. Veracruz: Naranjal, Near Cueva de Los Munecos, 700 m, 8 Apr 2005, S. Avendano & G. Alducin 5706 (holotype: XAL! [XAL 0005416, XAL 0005418]; isotypes: HEM [n. v.], MO!).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3E10BF94F81559809A078DF5A3A280B1.taxon	description	Description. Stem 10 - 40 cm long, 10 - 25 cm in diameter, epigeous, erect and decumbent. Cataphylls 1.5 - 3 x 2 - 4.2 cm wide at the base, persistent, triangular, densely tomentose at emergence, reddish brown and partially tomentose at maturity, apex acuminate. Leaves 2 - 7, 80 - 190 cm long, descending, reddish brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole 40 - 100 cm long, terete, linear, greenish brown in mature leaves; with 2 - 22 (28) thin prickles, 0.02 - 0.23 cm long. Rachis 40 - 123 cm long, terete, linear, brown and green in mature leaves, with prickles and occasionally unarmed. Leaflets 8 - 24 pairs, opposite to subopposite, insertion in one plane, oblong, in general longitudinally planar, not basally falcate, coriaceous, flat, green, adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical to asymmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 23 - 47.5 x 2.8 - 5 cm, 2.6 - 6.5 cm between leaflets; with articulations 0.7 - 1.2 cm wide, brown. Pollen strobili 20 - 23 cm long, 3.8 - 4.5 cm in diameter, solitary, cylindrical, erect, greenish yellow at emergence with reddish brown trichomes at maturity; peduncle 8 - 10.8 cm long, 1.2 - 1.6 cm in diameter, reddish brown to brown pubescent; microsporophylls 1 - 1.9 x 1 - 1.4 cm, obconic with a non-recurved distal face and lobate fertile portion, infertile portion 0.59 - 0.65 cm long and linear with straight horns 0.27 - 0.33 cm long, 0.67 - 0.80 cm and an acute angle between the horns. Ovulate strobili 9 - 11 cm long, 7 - 8 cm in diameter, solitary, cylindrical, erect, wine at emergence, wine with blackish brown trichomes at maturity, with an acute apex; peduncle 3 - 4 cm long, 1 - 1.2 cm in diameter, erect, brown pubescent; megasporophylls 18 - 49, 6 - 7 orthostichies with 3 - 7 sporophylls per orthostichy, 2.3 - 2.5 x 2 - 3 cm, with truncate distal face, horns straight and thin and 0.45 - 0.58 cm long, 0.99 - 1.48 cm between horns with a right angle between the horns. Seeds 1.2 - 2 cm long, 1.2 - 1.5 cm in diameter, ovate, sarcotesta whitish red when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3E10BF94F81559809A078DF5A3A280B1.taxon	distribution	Distribution and habitat. Ceratozamia decumbens is endemic to a small central mountain range in Veracruz State, Mexico (Fig. 16 C) growing on karstic rocks in mountain tropical forest and cloud forest at 450 - 1,100 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3E10BF94F81559809A078DF5A3A280B1.taxon	etymology	Etymology. The epithet alludes to the decumbent nature of trunks in older mature plants.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3E10BF94F81559809A078DF5A3A280B1.taxon	discussion	Discussion. Ceratozamia decumbens is distinguished from its most similar species (C. mexicana and C. morettii) by the ovulate strobili which are are wine red with blackish brown trichomes at maturity and an acute apex, whereas in C. mexicana they are green with blackish brown and gray trichomes with an acuminate apex, and C. morettii are green with with blackish brown trichomes with an apiculate apex. In terms of vegetative morphology, it differs from these species by its reddish brown leaves with whitish gray trichomes at emergence.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
3E10BF94F81559809A078DF5A3A280B1.taxon	materials_examined	Specimens examined. Mexico. Veracruz: Mun. Atoyac, 900 m, 28 Jan 1986, R. Acevedo R. 728 (XAL). Mun. Coetzala, 650 m, 30 Nov 2001, A. Rincon G. 2798 (MEXU, XAL); 870 m, 15 Jul 2015, L. Martinez-Dominguez et al. 655 - 683 (CIB). Mun. Cordoba, 1,100 m, 10 Jun 1985, A. Espiritu & J. L. Martinez 94 (XAL). Mun. Ixtaczoquitlan, 1,090 m, 25 May 1985, A. Perez P. 282 (XAL). Mun. Naranjal, 11 Sep 1982, A. P. Vovides 751 (XAL); 10 Oct 1993, Brigada T. Walters s / n (XAL); 11 Sep 1982, J. Rees 1690 (XAL); 10 Oct 1993, T. W. Walters 41277, 41308, 41397 (XAL). Mun. Tequila, 445 m, 28 Oct 2007, J. E. Rivera Hdez. & A. Vergara V. 4195 (MEXU, XAL); 959 m, 15 Jul 2015, F. Nicolalde-Morejon et al. 2259, 2260 (CIB); 959 m, 15 Jul 2015, L. Martinez-Dominguez et al. 684 - 703 (CIB). Mun. Tezonapa, 1 Dec 1995, M. A. Garcia B. 980 (XAL); 475 m, 24 Jun 1986, R. Robles G. 882 (XAL). Mun. Zongolica, 11 Mar 2011, L. Hermann Bojorquez G. et al. 2337 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
DAFB9C6F463C540F9EF531F67394F01C.taxon	description	Figs 2 A, B, 10 A, B, 15 H	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
DAFB9C6F463C540F9EF531F67394F01C.taxon	materials_examined	Type. Mexico. Veracruz: Mun. Atzalan, road Atzalan- Tlapacoyan, 3 km NE from Atzalan, 20 Jan 2012, M. Va ́ zquez-Torres & C. Carvajal-Herna ́ ndez 10200 ♀ (holotype: CIB! [acc. # 13915 UV]; isotypes: XAL [n. v.], XALU!).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
DAFB9C6F463C540F9EF531F67394F01C.taxon	description	Description. Stem 20 - 90 cm long, 25 - 40 cm in diameter, epigeous, erect and decumbent. Cataphylls 2 - 5.5 x 2.5 - 4.5 cm wide at the base, persistent, narrowly triangular, reddish brown, densely tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 10 - 100, 106 - 223 cm long, ascending, yellowish green at emergence with brown trichomes, glabrous at maturity. Petiole 30 - 87 cm long, terete, linear, light green at mature leaves; with 35 - 76 thin prickles, 0.21 - 0.76 cm long. Rachis 60 - 150 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 20 - 43 pairs, opposite to subopposite, insertion in one plane, lanceolate and oblong, longitudinally curved abaxially to planar, basally falcate to non-basally falcate, papyraceous to coriaceous, flat, green, adaxial side glaucous and glabrous and abaxial side glaucous, distal end with entire margins, acuminate and symmetrical to asymmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 22 - 45 x 2.3 - 4.6 cm, 1.5 - 5 cm between leaflets; articulations 0.6 - 1.6 cm wide, green. Pollen strobili 24 - 32 cm long, 5.5 - 7.6 cm in diameter, solitary, cylindrical, erect, greenish yellow at emergence, greenish yellow with blackish trichomes at maturity; peduncle 3.5 - 12.5 cm long, 1.3 - 2 cm in diameter, tomentose, reddish brown to light-brown; microsporophylls 1.5 - 2.5 x 1.3 - 2 cm, discoid and obconic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.30 - 0.45 cm long and orbicular with straight horns 0.15 - 0.25 cm long, 0.40 - 0.60 cm and a right angle between the horns. Ovulate strobili 17 - 40 cm long, 10 - 13.5 cm in diameter, solitary, cylindrical and globose, erect, dark green with blackish trichomes at emergence, green, generally glabrous at maturity, acute apex; peduncle 5.2 - 15 cm long, 1.8 - 2.2 cm in diameter, erect and pendulous, tomentose, brown to reddish brown; megasporophylls 48 - 266, 7 - 14 orthostichies with 6 - 19 sporophylls per orthostichy, 2.2 - 4.2 x 3.4 - 4.5 cm, with a truncate distal face, horns straight and 0.45 - 0.72 cm long, 1.05 - 1.67 cm between horns with a right angle between the horns. Seeds 2.0 - 3 cm long, 1.35 - 2.1 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
DAFB9C6F463C540F9EF531F67394F01C.taxon	distribution	Distribution and habitat. Ceratozamia delucana occurs in Veracruz and Puebla States, Mexico (Fig. 16 D) and grows on karstic rocks in evergreen tropical forest at 500 - 1,650 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
DAFB9C6F463C540F9EF531F67394F01C.taxon	etymology	Etymology. The epithet is in honor of Dr. Paolo De Luca, Professor at University of Naples Federico II and a researcher into the biology of Mexican cycads (Vazquez-Torres et al. 2013).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
DAFB9C6F463C540F9EF531F67394F01C.taxon	discussion	Discussion. Ceratozamia delucana is highly variable and in vegetative morphology similar to C. morettii. However, there are clear differences in their ovulate strobili. In C. delucana, ovulate strobili are green and generally without trichomes at maturity and have an acute apex, whereas in C. morettii they are green with blackish trichomes at maturity and have an apiculate apex. Additionally, C. delucana is a larger plant than C. morettii, with C. delucana having leaves up to 223 cm long with up to 43 pairs of leaflets and ovulate strobili 17 - 40 cm long.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
DAFB9C6F463C540F9EF531F67394F01C.taxon	materials_examined	Specimens examined. Mexico. Puebla: Mun. Hueytamalco, 520 m, 25 Feb 2008, G. Ibarra Manriquez et al. 5485 (MEXU, MO). Mun. Xochitlan de Vicente Suarez, 1,644 m, 9 Jun 2015, F. Nicolalde-Morejon & L. Martinez-Dominguez 2244, 2245 (CIB); 850 m, 25 Apr 1991, G. Villalobos & E. Guerrero C. 325 (MEXU); 1,644 m, 9 Jun 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 587 - 616 (CIB). Veracruz: Mun. Atzalan, 1,400 m, 20 Feb 2013, F. Nicolalde-Morejon et al. 1740 - 1742 (CIB); 1,400 m, 16 Aug 2014, F. Nicolalde-Morejon et al. 2125 - 2144 (CIB); 1,400 m, 13 Mar 2015, F. Nicolalde-Morejon et al. 2168 (CIB); 1,400 m, 27 Oct 2008, L. Lagunes-Galindo et al. 153 (CIB); 1396 m, 28 Oct 2008, L. Lagunes-Galindo et al. 155 (CIB); 1,400 m, 16 Aug 2014, L. Martinez-Dominguez et al. 163 (CIB); 863 m, 13 Mar 2015, L. Martinez-Dominguez et al. 228 - 238 (CIB); 1,400 m, 13 Mar 2015, L. Martinez-Dominguez et al. 239 - 248 (CIB). Mun. Las Minas, 2 Jun 1979, A. P. Vovides 427 (XAL); 1,500 m, 22 Sep 1988, C. Duran et al. 658 (XAL), 660 (MEXU, XAL); 1,500 m, 22 Sep 1988, C. Duran E. 659 (MEXU, XAL); 1,586 m, 20 Feb 2013, F. Nicolalde-Morejon et al. 1735 - 1739 (CIB); 1,621 m, 16 Aug 2014, F. Nicolalde-Morejon & L. Martinez-Dominguez 2107 - 2124 (CIB); 1,621 m, 16 Aug 2014, L. Martinez-Dominguez et al. 162 (CIB); 1,621 m, 18 Mar 2015, L. Martinez-Dominguez et al. 249 - 260 (CIB); 1 Mar 2005, 1,430 m, L. H. Bojorquez-Galvan 1349 (CIB); 1,420 m, 16 Mar 2005, L. H. Bojorquez-Galvan 1374 (CIB); 1,470 m, 28 Apr 2009, M. Vazquez-Torres et al. 8972 (CIB). Mun. Tlapacoyan, 900 m, 10 Jun 1970, Nevling & A. Gomez-Pompa 1083 (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
BA3DBA66D6785AABBC578BC7DE0749FD.taxon	description	Fig. 15 I	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
BA3DBA66D6785AABBC578BC7DE0749FD.taxon	materials_examined	Type. Mexico. Veracruz: Mun. Minatitlan, 21 Jun 1984, M. Vazquez-Torres 2842 ♀ (holotype: NY! [acc. # 1157 - 1166]; isotypes: CHAPA [n. v.], NY! [♂ acc. # 00001167 - 00001173], XALU [n. v.]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
BA3DBA66D6785AABBC578BC7DE0749FD.taxon	description	Description. Stem 25 - 50 cm long, 8 - 15.3 cm in diameter, semi-hypogeous, erect and decumbent. Cataphylls 4.5 - 7 x 4.2 - 7.5 cm wide at the base, persistent, triangular, reddish brown, densely tomentose at emergence, glabrous at maturity, apex acute. Leaves 2 - 22, 95 - 337.5 cm long, ascending, light green, glaucous at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 40 - 193 cm long, terete, linear, green to yellowish (pink in new leaves); with 8 - 35 thin prickles, 0.13 - 0.50 cm long. Rachis 53 - 218.5 cm long, terete, linear, green to yellowish in mature leaves, with prickles. Leaflets 6 - 17 pairs, opposite to subopposite, insertion in one plane, broadly obovate, in general longitudinally planar, not basally falcate, membranaceous, flat, green with adaxial and abaxial sides glabrous, distal end with sinuate margins, acuminate and asymmetrical at the apex, attenuate at base, with prominent and light-green veins; median leaflets 19.9 - 35.7 x 8.5 - 17.6 cm, 9 - 22 cm between leaflets; articulations 0.9 - 1.4 cm wide, yellow and green. Pollen strobili 27 - 35 cm long, 3 - 4.3 cm in diameter, solitary, cylindrical, erect, greenish yellow with reddish trichomes at emergence, greenish with black trichomes at maturity; peduncle 6 - 12 cm long, 1.5 - 2 cm in diameter, tomentose, reddish brown to brown; microsporophylls 1.0 - 1.5 x 0.5 - 1.3 cm, discoid with a non-recurved distal face and a lobate fertile portion, infertile portion 0.37 - 0.45 cm long and orbicular and rounded with straight horns 0.35 - 0.50 cm long, 0.60 - 0.72 cm and an acute angle between the horns. Ovulate strobili 15 - 20 cm long, 5 - 6 cm in diameter, solitary, cylindrical, erect, yellowish green to green with abundant, deep red trichomes at emergence, greenish brown with abundant, dark reddish brown trichomes at maturity, acuminate apex; peduncle 5 - 11.5 cm long, 1 - 2.5 cm in diameter, erect, tomentose, light brown; megasporophylls 35 - 64, 1.5 - 3 x 1.2 - 1.8 cm, 7 - 8 orthostichies with 5 - 8 sporophylls per orthostichy, 2.0 - 4.0 x 2.8 - 3.4 cm wide, with a truncate distal face, horns straight and 0.60 - 0.80 cm long, 0.99 - 1.40 cm between horns with an acute angle between the horns. Seeds 2.3 - 2.5 cm long, 1.5 - 1.7 cm in diameter, ovate, sarcotesta whitish red when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
BA3DBA66D6785AABBC578BC7DE0749FD.taxon	distribution	Distribution and habit. Ceratozamia euryphyllidia is endemic to the forest of Uxpanapa-Chimalapas in Oaxaca and Veracruz States, Mexico (Fig. 17 A) where it inhabits evergreen tropical rain forest on clay soils at the top and sides of hills between 100 and 630 m elevation.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
BA3DBA66D6785AABBC578BC7DE0749FD.taxon	etymology	Etymology. The specific epithet represents the very wide, diagnostic leaflets of this species. It comes from the Greek euryphyllos, which means, " wide leaflet ".	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
BA3DBA66D6785AABBC578BC7DE0749FD.taxon	discussion	Discussion. Ceratozamia euryphyllidia is easily diagnosable from its congeners in Mexico by its broadly obovate, lustrous and membranaceous leaflets asymmetrical apex of leaflets. Also, the ovulate strobili are greenish brown with an acuminate apex, and its megasporophylls are green with abundant dark brown trichomes. This species is most similar to C. hondurensis, but it can be distinguished by its sinuate margins at distal end of leaflets, whereas in C. hondurensis they are entire.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
BA3DBA66D6785AABBC578BC7DE0749FD.taxon	materials_examined	Specimens examined. Mexico. Veracruz: Mun. Minatitlan, 100 m, 24 Jul 1986, A. P. Vovides & E. R. Acosta 1108 (MO, XAL); 21 Jun 1984, M. Va ́ zquez-Torres 1157 (NY). Mun. Jesu ́ s Carranza, 24 May 1982, M. Va ́ zquez-Torres 2532 (CIB, MEXU); 26 Mar 1982, M. Va ́ zquez-Torres et al. 2374 (CHAPA, MEXU, NY); 100 m, 5 Jun 1989, M. Va ́ zquez-Torres & J. P. Sclavo 4037 (CIB); 18 Apr 1982, M. Va ́ zquez-Torres 2451 (CHAPA, MEXU, MO, NY); 120 m, 28 Jul 1983, M. Va ́ zquez-Torres 2614 (CIB, NY); 21 Jun 1984, M. Va ́ zquez-Torres 4126 (NY); 120 m, 28 Dec 1984, T. Wenth et al. 4732 (CHAPA). Mun. Uxpanapa, 398 m, 18 Mar 2014, F. Nicolalde-Morejo ́ n & L. Marti ́ nez-Domi ́ nguez 1984 (CIB); 19 Mar 2014, L. Marti ́ nez-Domi ́ nguez & F. Nicolalde-Morejo ́ n 71 - 77 (CIB). Mun. Hidalgotitla ́ n (plant cultivated at JBC-INECOL), V. E. Luna M. 1520 (XAL). Oaxaca: Mun. Santa Mari ́ a Chimalapa, 232 m, 13 May 1995, E. Torres B. 687 (SERO, XAL); 398 m, 18 Mar 2014, F. Nicolalde-Morejo ́ n & L. Martinez-Dominguez 1982, 1983 (CIB); 335 m, 5 May 1996, J. E. Rivera H. & S. Escobedo 102 (XAL); 630 m, 16 May 1998, J. E. Rivera H. et al. 700 (MEXU); 475 m, 21 Mar 1999, J. E. Rivera H. et al. 1195 (SERO); 398 m, 18 Mar 2014, L. Martinez-Dominguez & F. Nicolalde-Morejo ́ n 51 - 70 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
39A45C843C3E50ACBCF29799D3EB3B28.taxon	description	Figs 3 A, 9 C, 15 J	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
39A45C843C3E50ACBCF29799D3EB3B28.taxon	materials_examined	Type. Cultivated at Glasnevin, Ireland " Hort. Bot. Glasnevin ", 21 Mar 1878 (accessioned 1881), D. Moore s. n. (neotype, designated by Calonje and Sennikov 2017, pg. 161: K! [K 000501714, K 000501712, K 000501713]). Mexico. Hidalgo: Mun. Molango, km 3 carretera Molango-Xochicoatlan, 1,860 m, 31 Mar 2015, L. Marti ́ nez-Domi ́ nguez et al. 493 ♀ (epitype, designated by Martinez-Dominguez et al. 2018 a: 105: CIB! [acc. # 17465 UV]; isoepitype: MEXU! [acc. # 1520508, 1520282, 1520519]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
39A45C843C3E50ACBCF29799D3EB3B28.taxon	description	Description. Stem 20 - 90 cm long, 25 - 40 cm in diameter, epigeous, erect and decumbent. Cataphylls 2 - 4.5 x 2 - 3.5 cm wide at the base, persistent, narrowly triangular, reddish brown, densely brownish tomentose at emergence, tomentose at maturity, apex acuminate. Leaves 10 - 70, 92 - 215 cm long, ascending, light green and dark brown at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 40 - 95 cm long, terete, linear, dark green in mature leaves; with 14 - 60 (66) thin prickles, 0.09 - 0.33 cm long. Rachis 65 - 150 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 28 - 67 pairs, opposite to subopposite, insertion in one plane, lanceolate, abaxially curved longitudinally, basally falcate, papyraceous, flat, green, with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and green-light brown veins; median leaflets 16.6 - 42 x 1.3 - 2.1 cm, 0.6 - 2 cm between leaflets; articulations 0.6 - 1.3 cm wide, green and brown. Pollen strobili 22 - 30 cm long, 5 - 8 cm in diameter, solitary, cylindrical, erect, brownish yellow at emergence, greenish brown with reddish brown trichomes at maturity; peduncle 5 - 14.5 cm long, 1.6 - 2.3 cm in diameter, tomentose, reddish brown to brown; microsporophylls 1.6 - 2.4 x 1.2 - 1.6 cm, obconic with a non-recurved distal face and a deeply lobate fertile portion, infertile portion 0.45 - 0.56 cm long and orbicular with straight horns 0.16 - 0.29 cm long, 0.53 - 0.80 cm and a right angle between the horns. Ovulate strobili 24 - 35 cm long, 8.5 - 15 cm in diameter, solitary, cylindrical, erect, green with brown trichomes at emergence, brown-green with dark brown trichomes at maturity, acuminate apex; peduncle 4 - 15.5 cm long, 1.8 - 2.5 cm in diameter, erect or pendulous, tomentose, brown to reddish brown; megasporophylls 49 - 195, 7 - 15 orthostichies with 7 - 15 sporophylls per orthostichy, 2.3 - 3.5 x 3.0 - 4.0 cm, with a truncate distal face, horns straight and 0.70 - 0.99 cm long, 1.7 - 2.1 cm between horns and an obtuse angle between the horns. Seeds 2 - 2.6 cm long, 1.5 - 2 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
39A45C843C3E50ACBCF29799D3EB3B28.taxon	distribution	Distribution and habitat. Ceratozamia fuscoviridis is endemic to south to central Carso Huasteco in Mexico from Hidalgo to northwest of Veracruz States (Fig. 17 B) in cloud forests between 1,300 - 1,900 m elevation.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
39A45C843C3E50ACBCF29799D3EB3B28.taxon	etymology	Etymology. The epithet refers to the dark-brown color of the leaf at emergence.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
39A45C843C3E50ACBCF29799D3EB3B28.taxon	discussion	Discussion. Ceratozamia fuscoviridis is polymorphic within populations because individual plants may have either a light green or dark-brown leaf color at emergence. Individuals with dark-brown leaf at emergence also have a brownish abaxial side. This species is most geographically proximate to C. chamberlainii, but it can be distinguished by lanceolate and papyraceous leaflets, a petiole armed with long and thin prickles; and ovulate strobili that are brownish green with dark trichomes at maturity. Osborne et al. (2006) intended to validate Ceratozamia fuscoviridis D. Moore, which was invalidly published in 1878. However, in 2017, Calonje & Sennikov published the correction to this name using a brief description by William Bull in horticultural catalogues. This work is the valid publication for the specie and the name " C. fuscoviridis D. Moore " is an isonym with no nomenclatural status.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
39A45C843C3E50ACBCF29799D3EB3B28.taxon	materials_examined	Specimens examined. Mexico. Hidalgo: Mun. Eloxochitlan, 18 Mar 1995, I. Luna-Vega 54716 (FCME). Mun. Metztitlan, 30 Dec 1992, J. L. Lopez-Garcia 449 (ENCB, IBUG, MEXU). Mun. Molango de Escamilla, 1,380 m, 29 May 1999, A. P. Vovides 1298 (XAL); 1,400 m, 29 May 1999, A. P. Vovides 1301 (XAL); 1,500 m, 24 Jul 2008, A. Vite-Reyes et al. 6 (XAL); 1,860 m, 31 Mar 2015, F. Nicolalde-Morejon et al. 2209 - 2211 (CIB); 1,860 m, 31 Mar 2015, L. Martinez-Dominguez et al. 485 - 492, 494 - 514 (CIB); T. W. Walters 2001 - 03 - A (XAL). Mun. Tenango de Doria, 1,700 m, 12 Mar 1993, I. Luna-Vega 914 (FCME). Mun. Tlanchinol, 1,450 m, 25 Aug 1992, I. Luna-Vega s / n (FCME); 1,420 m, 7 Oct 1992, I. Luna-Vega 625 (FCME, XAL); 25 Aug 1992, I. Luna-Vega 789 (XAL); 1,312 m, 31 Mar 2015, F. Nicolalde-Morejon et al. 2204 - 2208 (CIB); 1,312 m, 31 Mar 2015, L. Martinez-Dominguez et al. 455 - 484 (CIB). Mun. Zacualtipan de Angeles, 23 Jan, 1983, J. Rees 389 (CHAPA, FCME, MEXU); 1,360 m, 4 Dec 1974, J. Rees 1611 (CHIP, XAL); 4 Dec 1974, J. Rees 6339 (IEB). Veracruz: Mun. Huayacocotla, 1,913 m, 23 Feb 2005, D. Saavedra Millan 64 (FCME); 1,850 m, 13 Mar 1980, J. Palma G. 63 (XAL); 1,844 m, 1 Apr 2015, F. Nicolalde-Morejon et al. 2212 - 2214 (CIB); 1,700 m, 24 Mar 1981, L. Ballesteros & F. Ballesteros 460 (XAL); 1,550 m, 23 Apr 1981, L. G. Juarez G. 47 (XAL); 1,844 m, 1 Apr 2015, L. Martinez-Dominguez et al. 515 - 544 (CIB); 1,900 m, 11 Feb 1972, R. Hernandez M. 1507 (MEXU, XAL); 26 Feb 1975, V. Sosa 59 (XAL).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2B539FB3369F5343B7C062BA3841291F.taxon	description	Fig. 18	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2B539FB3369F5343B7C062BA3841291F.taxon	materials_examined	Type. Cultivated in Baton Rouge, Louisiana USA at 5988 South Pollard Parkway (plants originally from several km N of Xilitla, San Luis Potosi, Mexico), G. Landry 76521 ♂ (holotype: GH! [00003274]; isotypes: FTG!, LSU! [LSU 00048484], MEXU! [MEXU 00443083, MEXU 00443084, MEXU 00443085], MICH! [1050284 A, 1050284 B], NY! [00001153 - 00001156], US! [00011993]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2B539FB3369F5343B7C062BA3841291F.taxon	description	Description. Stem 10 - 20 cm long, 10 - 15 cm in diameter, semi-hypogeous, erect. Cataphylls 2.1 - 4.2 x 0.8 - 1.9 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 2 - 7, 95 - 202 cm long, ascending, reddish brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole 43 - 89 cm long, terete, linear, greenish brown and green in adult leaves; with 2 - 12 thin prickles, 0.01 - 0.2 cm long. Rachis 60 - 130 cm long, terete, linear, greenish brown in mature leaves, with prickles and occasionally unarmed. Leaflets in 5 - 11 fascicles, 16 - 56 leaflets in total, clustered, insertion in one plane, oblong, in general longitudinally planar, basally falcate to non-basally falcate, membranaceous, flat, green with adaxial and abaxial sides glaucous, distal end with entire margins, acuminate and symmetrical to asymmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 14.5 - 24 x 2.4 - 5 cm, 6 - 15 cm between leaflets; articulations 0.2 - 0.5 cm wide, brown and green. Pollen strobili 8 - 12.5 cm long, 2 - 2.5 cm in diameter, solitary, cylindrical, erect, brown with reddish brown trichomes at emergence, reddish brown at maturity; peduncle 6 - 8 cm long, 0.8 - 1 cm in diameter, tomentose, reddish brown to brown; microsporophylls 0.8 - 1.2 x 0.6 - 1 cm, discoid with non-recurved distal face and a lobate fertile portion, infertile portion 0.23 - 0.29 cm long and rounded with curved horns, 0.20 - 0.25 cm long, 0.39 - 0.50 cm and an acute angle between the horns. Ovulate strobili 10 - 15 cm long, 6 - 9 cm in diameter, solitary, cylindrical, erect, green at emergence with brown trichomes, green with brown to blackish trichomes at maturity, acuminate apex; peduncle 7.5 - 16 cm long, 1.2 - 1.5 cm in diameter, erect, tomentose, brown; megasporophylls 20 - 56, 5 - 8 orthostichies with 4 - 7 sporophylls per orthostichy, 2 - 3.7 x 2 - 4 cm, with a prominent distal face, horns straight and 0.30 - 0.50 cm long, 1.70 - 1.90 cm between horns with a right angle between the horns. Seeds 1.3 - 2.3 cm long, 1.2 - 1.5 cm in diameter, ovate, sarcotesta whitish red when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2B539FB3369F5343B7C062BA3841291F.taxon	distribution	Distribution and habitat. Ceratozamia hildae is endemic to Sierra Gorda in Mexico, particularly in San Luis Potosi ́ and Queretaro (Fig. 17 C), where it occurs in the evergreen tropical forests on karstic rocks at 300 - 1,200 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2B539FB3369F5343B7C062BA3841291F.taxon	etymology	Etymology. The epithet is in honor of Hilda Guerra Walker, daughter of the original collector (Luciano E. Guerra, plant collector from Mission Texas) (Landry and Wilson 1979).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2B539FB3369F5343B7C062BA3841291F.taxon	discussion	Discussion. Ceratozamia hildae is easily distinguished from other members of the group by its clustered membranous oblong leaflets (Fig. 18 A).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2B539FB3369F5343B7C062BA3841291F.taxon	materials_examined	Specimens examined. Mexico. Queretaro: Mun. Arroyo Seco, 1,181 m, 9 Nov 2003, J. A. Perez de la Rosa & G. Vargas A. 1908 (IBUG). Mun. Jalpan de Serra, 850 m, 29 Dec 1977, A. P. Vovides 337 (XAL); 29 Dec 1977, A. P. Vovides s / n (IEB); 1,200 m, 8 Mar 1991, B. Servin 870 (MEXU); 1,200 m, 20 Mar 1991, B. Servin 986 (MEXU); 8 Mar 1991, B. Servin 6328 (IEB); 20 Apr 1991, B. Servin 6330 (IEB); Sep 1977, J. Rees s / n (IEB); 312 (XAL). San Luis Potosi: Mun. Aquismon, 300 m, 22 Sep 1977, A. P. Vovides & J. Rees 312 (MEXU, XAL); 357 m, 20 Mar 2016, F. Nicolalde-Morejon et al. 2391 - 2406 (CIB); 357 m, 20 Mar 2016, L. Martinez-Dominguez et al. 910 - 921, 923, 940 (CIB), 922 (CIB, MEXU); 26 Nov 1970, S. Longoria s / n (MEXU; US); 617 m, 15 Jan 2001, T. W. Walters TW- 2001 - 14 - A (MEXU, XAL), TW- 2001 - 14 - B (XAL), TW- 2001 - 14 - C (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5541FFD41AE65C36A5BC8A46D45871C4.taxon	description	Fig. 15 K	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5541FFD41AE65C36A5BC8A46D45871C4.taxon	materials_examined	Type. HONDURAS. Atlantida: Mun. Esparta, 0.5 km SE of Jilamo Nuevo along the Rio Jilamito, 13 Apr 1994, A. E. Brand & R. Zuniga 2830, (lectotype, designated here: MO! [acc. # 5943287 - 5943289]; isolectotype: MEXU! [MEXU 01347996 - MEXU 01347999]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5541FFD41AE65C36A5BC8A46D45871C4.taxon	description	Description. Stems 20 - 50 cm long, 20 cm in diameter, semi-hypogeous, erect and decumbent. Cataphylls 4.5 - 7 x 4.5 - 7.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 3 - 25, 120 - 322 cm long, ascending, light green at emergence with whitish gray trichomes, glabrous at maturity. Petiole 56 - 150 cm long, terete, linear, green in mature leaves; with 8 - 34 thin prickles, 0.26 - 0.50 cm long. Rachis 80 - 200 cm long, terete, linear, with prickles, green in mature leaves. Leaflets 10 - 18 pairs, opposite to subopposite, insertion in one plane, obovate to broadly oblanceolate, in general longitudinally planar, generally not basally falcate, membranaceous, flat, green with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with prominent and light-green veins; median leaflets 23 - 34.5 x 8 - 12.8 cm, 8.5 - 16 cm between leaflets; articulations 0.5 - 1.6 cm wide, green and yellow. Pollen strobili 30 - 35 cm long, 4 - 5 cm in diameter, solitary, cylindrical, erect, yellowish green at emergence, yellowish cream with blackish trichomes at maturity; peduncle 2 - 4 cm long, 2 - 3 cm in diameter, tomentose, brown; microsporophylls 0.9 - 1.2 x 0.7 - 1.2 cm, discoid with a non-recurved distal face and lobate fertile portion, infertile portion 0.52 - 0.60 cm long, and orbicular and rounded with straight horns 0.18 - 0.23 cm long, 0.50 - 0.62 cm and an acute angle between the horns. Ovulate strobili 18 - 22 cm long, 8 - 10 cm in diameter, solitary, cylindrical, erect, yellowish green to green with abundant, deep red trichomes at emergence, greenish brown with abundant, blackish trichomes at maturity, mucronate apex; peduncle 5 - 15 cm long, 1.0 - 3.0 cm in diameter, erect, tomentose, brown; megasporophylls 72 - 120, 9 - 11 orthostichies with 8 - 11 sporophylls per orthostichy, 1.5 - 2.5 x 4 - 5.5 cm, with a prominent distal face, horns curved and 0.60 - 0.80 cm long, 0.99 - 1.40 cm between the horns with an acute angle between horns. Seeds 2.1 - 2.6 cm long, 1.46 - 1.89 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5541FFD41AE65C36A5BC8A46D45871C4.taxon	distribution	Distribution and habitat. Ceratozamia hondurensis is endemic to Honduras in the Atlantida department (Fig. 17 D), where it grows in evergreen tropical forest between 20 - 600 m elevation.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5541FFD41AE65C36A5BC8A46D45871C4.taxon	etymology	Etymology. The specific epithet alludes to the endemism of the species in Honduras.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5541FFD41AE65C36A5BC8A46D45871C4.taxon	discussion	Discussion. Ceratozamia hondurensis is similar in leaf morphology to C. euryphyllidia. It differs in its leaflets with a symmetric apex and an entire margin at distal end as compared to the asymmetric apex and sinuate margin at distal end of the leaflets in C. euryphyllidia. Additionally, C. hondurensis differs in reproductive structures; the ovulate strobilus has mucronate apex, whereas in C. euryphyllidia the ovulate strobilus has long (up to 5 cm) acuminate apex. In protologue, two specimens from " A. E. Brand & R. Zuniga 2830 " were cited as types: holotype in EAP and isotype in MO. The type specimen was not deposited in EAP (the herbarium has no record of this material nor was it found in unprocessed material). In addition, we consulted TEFH, the other herbarium in Honduras, with the same result. However, we found a duplicate specimen in MEXU. Thus, we are here designating the specimen in MO cited as isotype as the lectotype and the specimen at MEXU as the isolectotype.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5541FFD41AE65C36A5BC8A46D45871C4.taxon	materials_examined	Specimens examined. Honduras. Atlantida: Mun. La Ceiba, 225 m, 14 Apr 1996, D. R. Hodel & Schleder 1485 (MO); 217 m, 29 Jul 2003, J. Haynes et al. 47 (TEFH); 151 m, 19 May 2022, L. Martinez-Dominguez et al. 2440, 2443, 2444, 2446 - 2448 (TEFH). Mun. Tela, 0 - 500 m, 10 Feb 1994, C. Nelson et al. 17586 (TEFH); 200 - 500 m, 9 Apr 1994, D. L. Hazlett et al. 8036 (MO); 615 m, 13 Jul 2008, G. Sandoval et al. 1312 (TEFH); 615 m, 31 Jul 2003, J. Haynes et al. 40 (TEFH); Cultivated, 615 m, 16 Mar 2017, L. Ferrufino et al. 777 (TEFH); 163 m, 17 May 2022, L. Martinez-Dominguez et al. 2421, 2423, 2425, 2428 - 2430 (TEFH).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9FBAF21B3BD95E2C88E4645B177BC4CB.taxon	description	Fig. 15 L	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9FBAF21B3BD95E2C88E4645B177BC4CB.taxon	materials_examined	Type. Mexico. Veracruz: Mun. Tepetzintla, Sierra de San Juan Otontepec, 1,300 m, 16 Dec 1981, G. Castillo-Campos et al. 2567 ♀ (holotype: XAL! [XAL 0016937]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9FBAF21B3BD95E2C88E4645B177BC4CB.taxon	description	Description. Stem 20 - 50 cm long, 14 - 20 cm in diameter, semi-hypogeous, erect and decumbent. Cataphylls 3.0 - 4.0 x 2.5 - 3.0 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 4 - 7, 50 - 80 cm long, ascending, light green at emergence with whitish gray trichomes, glabrous at maturity. Petiole 20 - 40 cm long, terete, linear, green in mature leaves; with 10 - 30 thin prickles 0.21 - 0.35 cm long. Rachis 40 - 70 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 8 - 18 pairs, opposite to subopposite, insertion in one plane, oblong, in general longitudinally planar, not basally falcate, coriaceous, flat, green with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acuminate and asymmetrical at the apex, attenuate at base, with prominent and light-green veins; median leaflets 12 - 22 x 2.5 - 6 cm, 1.6 - 5 cm between leaflets; articulations 0.4 - 1.1 cm wide, green. Pollen strobili 15 - 18 cm long, 2.5 - 3.2 cm in diameter, solitary, cylindrical, erect, yellowish green at emergence, yellowish cream with blackish trichomes at maturity; peduncle 6 - 23 cm long, 1.6 - 2.2 cm in diameter, tomentose, brown; microsporophylls 0.8 - 1.2 x 0.72 - 1.0 cm, discoid with a non-recurved distal face and a lobate fertile portion, infertile portion 0.28 - 0.35 cm long and orbicular with straight horns 0.11 - 0.18 cm long, 0.50 - 0.68 cm and an acute angle between the horns. Ovulate strobili 13.5 - 18 cm long, 6 - 10 cm in diameter, solitary, cylindrical, erect, green at emergence, dark green with abundant blackish trichomes at maturity, acuminate apex; peduncle 3 - 8 cm long, 1.0 - 2.2 cm in diameter, erect, tomentose, brown; megasporophylls 54 - 99, 9 - 10 orthostichies with 6 - 9 sporophylls per orthostichy, 1.4 - 1.8 x 1.8 - 2.0 cm, with a truncate distal face, horns straight and 0.40 - 0.56 cm long, 1.0 - 1.60 cm between the horns with an acute angle between the horns. Seeds 1.1 - 1.5 cm long, 1.0 - 1.2 cm in diameter, ovate, sarcotesta light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9FBAF21B3BD95E2C88E4645B177BC4CB.taxon	distribution	Distribution and habitat. Ceratozamia huastecorum is endemic to the Sierra de Otontepec in the northwest of Veracruz State, Mexico (Fig. 19 A) where it occurs in cloud forest on clay soils with rocky outcrops at 800 to 1,300 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9FBAF21B3BD95E2C88E4645B177BC4CB.taxon	etymology	Etymology. The specific epithet commemorates the Huasteca region, where this species is found; this region has cultural importance in Veracruz, Mexico.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9FBAF21B3BD95E2C88E4645B177BC4CB.taxon	discussion	Discussion. Ceratozamia huastecorum has a close morphological similarity to C. latifolia. However, C. huastecorum has light-green emergent leaves and coriaceous leaflets, whereas in C. latifolia the leaflets are reddish brown at emergence and papyraceous. In addition, the ovulate strobili are larger than C. latifolia, with 9 to 10 orthostichies as compared to C. latifolia with 4 - 7 orthostichies.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9FBAF21B3BD95E2C88E4645B177BC4CB.taxon	materials_examined	Specimens examined. Mexico. Veracruz: Mun. Tepetzintla, 1,250 m, 26 Aug 1981, G. Castillo-Campos & A. Benavides M. 2215 (XAL); 1,200 m, 27 Aug 1981, G. Castillo-Campos & A. Benavides M. 2232, 2244 (XAL); 820 m, 14 Dec 1981, G. Castillo-Campos 2480, 2481, 2482 (XAL); 850 m, 10 Jan 1986, G. Castillo-Campos 4472 (XAL); 20 Sep 1989, P. Zamora C. et al. 1197 (CH, MEXU, XAL); 21 Sep 1989, P. Zamora et al. 1249 (MEXU, XAL).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AD3592D891D25D3CB414A71FEFB90639.taxon	description	Figs 3 D, 7 A, 9 B	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AD3592D891D25D3CB414A71FEFB90639.taxon	materials_examined	Type. Cultivated in St. Petersburg, Russia " Ex Horto Petropolitano ", 1856, E. Regel s. n. (holotype: LE! [LE 00009046]; isotype: U! [U 0007272]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AD3592D891D25D3CB414A71FEFB90639.taxon	description	Description. Stem 10 - 30 cm long, 10 - 25 cm in diameter, semi-hypogeous, erect. Cataphylls 1.5 - 4 x 2 - 3.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 1 - 11, 80 - 133 cm long, ascending, reddish brown at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 30 - 72 cm long, terete, linear, greenish brown in mature leaves; with 1 - 10 thin prickles, 0.01 - 0.15 cm long. Rachis 40 - 72 cm long, terete, linear, greenish brown in mature leaves, with prickles and rarely unarmed. Leaflets 22 - 50 pairs, opposite to subopposite, insertion in one plane, linear, adaxially curved, basally falcate, papyraceous, caniculate, green, adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and light veins; median leaflets 17 - 32 x 0.6 - 1 cm, 0.3 - 2.5 cm between leaflets; articulations 0.2 - 0.8 cm wide, brown. Pollen strobili 11 - 15 cm long, 1.6 - 2.5 cm in diameter, solitary, cylindrical, erect, greenish yellow with brown trichomes at emergence, yellowish brown with reddish brown trichomes at maturity; peduncle 8 - 14 cm long, 0.7 - 1 cm in diameter, reddish brown to brown pubescent; microsporophylls 0.6 - 1.1 x 0.5 - 1 cm, discoid with a non-recurved distal face and a lobate fertile portion, infertile portion 0.28 - 0.32 cm long and linear with straight horns 0.05 - 0.25 cm long, 0.45 - 0.53 cm and an acute angle between the horns. Ovulate strobili 13 - 21 cm long, 7 - 9 cm in diameter, solitary, cylindrical, erect, green at emergence with brown trichomes, grayish light green with black trichomes at maturity, acuminate apex; peduncle 9 - 16 cm long, 1 - 2 cm in diameter, erect, tomentose, brown; megasporophylls 30 - 72, 6 - 8 orthostichies with 5 - 9 sporophylls per orthostichy, 2.5 - 4.0 x 2.3 - 3.5 cm, with a truncate distal face, horns straight and 0.30 - 0.52 cm long, 0.60 - 0.94 cm between horns with an obtuse angle between the horns. Seeds 1.5 - 2.3 cm long, 1.2 - 1.8 cm in diameter, ovate, sarcotesta whitish red when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AD3592D891D25D3CB414A71FEFB90639.taxon	distribution	Distribution and habitat. Ceratozamia kuesteriana is endemic to Tamaulipas State, Mexico (Fig. 19 B), where it occurs on karstic rocks in pine-oak and cloud forests at 1,100 - 1,500 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AD3592D891D25D3CB414A71FEFB90639.taxon	etymology	Etymology. The specific epithet is in honor of Baron K. von Kuester.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AD3592D891D25D3CB414A71FEFB90639.taxon	discussion	Discussion. Ceratozamia kuesteriana is similar to C. sabatoi, but it is easily distinguished by its canaliculate leaflets and ascending leaves.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AD3592D891D25D3CB414A71FEFB90639.taxon	materials_examined	Specimens examined: Mexico. Tamaulipas: Mun. Gomez Farias, 1,260 m, 1 May 1967, A. Gomez-Pompa 2029 (MEXU); 1,150 m, 12 Aug 1983, A. P. Vovides 791, 800, 2056 (XAL); 13 Aug 1983, A. P. Vovides & G. Pattison 771, 772, 801 (XAL); 27 Jun 2017, C. I. Carvajal-Hernandez et al. 1355 (CIB); 1,200 - 1400 m, 20 Nov 1970, F. Gonzalez-Medrano & E. Martinez 3288 (MEXU); 1,100 m, 21 Jan 1970, F. Gonzalez-Medrano et al. 3362 (MEXU, MO); 13 Aug 1985, L. Trejo s / n (UAT); 1,050 m, 30 Mar 1961, P. S. Martin & C. Saravia 1170 (ENCB). Mun. Ocampo, 1,255 m, 14 Jan 2001, T. W. Walters TW- 201 - 11 - A (XAL), TW- 201 - 11 - B (MEXU). Mun. Tula, 1,450 m, 21 Jul 1983, D. W. Stevenson 569 H, 569 G, 569 K (MEXU); 1,296 m, 19 Mar 2016, F. Nicolalde-Morejon et al. 2357 - 2365 (CIB); 1,296 m, 19 Mar 2016, L. Martinez-Dominguez et al. 854 - 867, 869 (CIB), 868 (CIB, MEXU); 1,140 m, 19 Mar 2016, L. Martinez-Dominguez et al. 870 - 881 (CIB); 24 Apr 2001, S. Avendano 5328 (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
94684D81DE30568F817819575172F967.taxon	description	Figs 7 B, 10 A	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
94684D81DE30568F817819575172F967.taxon	materials_examined	Type. Mexico. San Luis Potosi: Route 70, 46 km West of Ciudad Valles, 650 m, 20 Jul 1983, D. E. Stevenson 565 E (neotype, designated by Stevenson and Sabato 1986, pg. 579: NY! [00001117]; isoneotypes: MEXU! [MEXU 00469173, MEXU 00469148]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
94684D81DE30568F817819575172F967.taxon	description	Description. Stem 5 - 15 cm long, 10 - 25 cm in diameter, semi-hypogeous, erect. Cataphylls 1.5 - 3 x 2 - 4 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 1 - 8, 53 - 163.5 cm long, descending, reddish brown at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 25 - 80 cm long, terete, linear, greenish brown in mature leaves, generally unarmed, rarely up to 3 prickles, 0.08 - 0.10 cm long. Rachis 25 - 110 cm long, terete, linear, greenish brown in mature leaves, unarmed. Leaflets 7 - 22 pairs, opposite to subopposite, insertion in one plane, oblong, in general longitudinally planar, basally falcate, papyraceous, flat, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and asymmetrical at the apex, attenuate at base, with conspicuous and indistinct veins; median leaflets 12 - 28 x 2.3 - 5.1 cm, 1.7 - 12.5 cm between leaflets; articulations 0.4 - 1.1 cm wide, brown. Pollen strobili 10 - 20 cm long, 2.1 - 2.5 cm in diameter, solitary, cylindrical, erect, greenish yellow with reddish brown trichomes at emergence, reddish brown at maturity; peduncle 5 - 15 cm long, 0.79 - 1.5 cm in diameter, reddish brown to brown pubescent; microsporophylls 0.5 - 1.3 x 0.5 - 1.1 cm, discoid with a non-recurved distal face and a lobate fertile portion, infertile portion 0.20 - 0.30 cm long and rounded with straight horns 0.1 - 0.20 cm long, 0.40 - 0.55 cm and a right angle between the horns. Ovulate strobili 6.5 - 16 cm long, 5.5 - 8.0 cm in diameter, solitary, globose, erect, light green at emergence with brown trichomes, greyish light brown with brown trichomes at maturity, apiculate apex; peduncle 4 - 13.5 cm long, 1.3 - 1.8 cm in diameter, erect, brown pubescent; megasporophylls 24 - 63, 5 - 7 orthostichies with 4 - 9 sporophylls per orthostichy, 1.0 - 2.7 x 1.6 - 2.9 cm, with a prominent distal face, horns straight and 0.36 - 0.76 cm long, 0.54 - 0.63 cm between horns with an obtuse angle between the horns. Seeds 1.5 - 2.2 cm long, 1.2 - 1.6 cm in diameter, ovate, sarcotesta whitish red when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
94684D81DE30568F817819575172F967.taxon	distribution	Distribution and habitat. Ceratozamia latifolia is distributed widely in the mountainous region of San Luis Potosi ́ State, Mexico (Fig. 19 C), where it occurs in pine-oak, oak and cloud forests between 600 - 1,100 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
94684D81DE30568F817819575172F967.taxon	etymology	Etymology. The specific epithet is derived from the Latin word " latus " (wide) and " folium " (leaves).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
94684D81DE30568F817819575172F967.taxon	discussion	Discussion. Ceratozamia latifolia differs from C. chamberlainii by its unarmed petiole or if armed with no more than 3 prickles, its papyraceous leaflets, and discoid microsporophylls with a rounded distal face and horns up to 0.20 cm long. Additionally, this species is characterized by reddish brown leaves at emergence and ovulate strobilus is greyish light brown with brown trichomes at maturity and an apiculate apex.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
94684D81DE30568F817819575172F967.taxon	materials_examined	Specimens examined. Mexico. San Luis Potosi: Cultivated, 24 Jan 1994, F. Garcia S. s / n (SLPM); Dec 1977, N. F. McCarten 2552 (ENCB). Mun. Alaquines, 1,428 m, 21 Dec 2008, J. Fortanelli M. & H. A. Castillo 141 (SLPM). Mun. Cardenas, 950 m, 12 to 15 Sep 1967, J. Rzedowski 24746 (ENCB). Mun. Ciudad del Maiz, 750 m, 13 Oct 1968, H. Puig 3420 (ENCB, P); 25 Apr 2001, S. Avendano 5320 (MEXU); 900 - 1,300 m, 5 Feb 1984, S. Sabato et al. 2340 (ENCB, MEXU, MO). Mun. El Naranjo, 7 Nov 1951, Clint 144 (US); 20 Jul 1997, D. W. Stevenson 567 (XAL); 900 m, 19 Mar 2016, F. Nicolalde-Morejon et al. 2375 - 2389 (CIB); 900 m, 19 Mar 2016, L. Martinez-Dominguez et al. 894 - 909 (CIB); 895 m, 13 Jan 2001, T. W. Walters TW- 2001 - 08; 895 M, 14 Jan 2001, TW- 2001 - 10 A, B (MEXU, XAL). Mun. Rayon, 812 m, 5 Aug 2003, A. P. Vovides et al. 1466 (XAL); 20 Jul 1983, D. W. Stevenson 1117 (NY); 650 m, 20 Jul 1983, D. W. Stevenson 565 A (MEXU), 565 B, C (MEXU, XAL); 993 m, 18. Mar 2016, F. Nicolalde-Morejon et al. 2320 - 2334 (CIB); 1,100 m, 30 Jun 1962, F. Medellin L. 1330 (ENCB, MEXU, SLPM); 1,221 m, 6 Apr 2013, H. A. Castillo-Gomez et al. 558 (SLPM); 600 m, 17 Jul 1988, J. A. Perez de la Rosa & L. M. Gonzalez-Villarreal 1564 (IBUG); 993 m, 18 Mar 2016, L. Martinez-Dominguez et al. 811 - 827 (CIB); 24 Apr 2001, S. Avendano 5282 (MEXU). Mun. Tamasopo, 700 m, 15 Aug 2003, A. P. Vovides et al. 1465 (MEXU, XAL); 2 Jun 1968, F. Medellin L. 27241 (ENCB); 2 Jun 1968, F. Medellin L. s / n (ENCB, IBUG, MEXU, SLPM); 716 m, 18. Mar 2016, F. Nicolalde-Morejon et al. 2335 - 2348 (CIB); 716 m, 18 Mar 2016, L. Martinez-Dominguez et al. 828 - 844 (CIB); 750 m, 11 Sep 1967, J. Rzedowski 24571 (ENCB); 900 m, 24 May 1981, P. A. Fryxell & W. R. Anderson 3586 (NY; US); 856 m, 19 Jan 2013, U. Pineda M. 70 (SLPM).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
20036E4799B65E3BA2EF245D63B9AD7C.taxon	description	Fig. 7 C	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
20036E4799B65E3BA2EF245D63B9AD7C.taxon	materials_examined	Type. Mexico. Guerrero: Mun. Tlacoachistlahuaca, 3 km NW de San Pedro Cuitlapan, 1,400 m, 26 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1867 ♀ (holotype: CIB! [acc. # 22405 UV]; isotypes: MEXU!, NY!).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
20036E4799B65E3BA2EF245D63B9AD7C.taxon	description	Description. Stem 30 - 150 cm long, 11 - 35 cm in diameter, epigeous, erect to decumbent. Cataphylls 9 - 11 x 2.5 - 3 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose abaxially at emergence, pubescent at maturity, apex acuminate. Leaves 7 - 37 (55), 93.5 - 281 cm long, descending, green at emergence with sparse reddish brown pubescent, glabrous at maturity. Petiole 45 - 85 cm long, terete, linear, copperish green in mature leaves; with 50 - 75 thin prickles, 0.48 - 0.68 cm long. Rachis 75 - 196 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 22 - 61 pairs, opposite to subopposite, insertion in one plane, linear, abaxially curved, not basally falcate, membranaceous, flat, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and greenish veins; median leaflets 28 - 43.5 x 1.9 - 2.8 cm, 1.8 - 2.8 cm between leaflets; articulations 0.70 - 1.15 cm, generally copperish green. Pollen strobili 40 - 45 cm long 6.0 - 7.8 cm in diameter, generally solitary (rarely 2), cylindrical, erect, brownish yellow at emergence, yellowish green with brownish trichomes at maturity; peduncle 13 - 19 cm long, 1.5 - 2.0 cm in diameter tomentose, reddish brown to brown; microsporophylls 2.1 - 2.45 x 1.09 - 1.30 cm, obconic with a non-recurved distal face and a deeply lobate fertile portion, infertile portion 0.83 - 0.96 cm long and linear with straight and thin horns and 0.1 - 0.23 cm long, 0.44 - 0.56 cm and an acute angle between the horns. Ovulate strobili 23.5 - 28 cm long, 9.5 - 11 cm in diameter, solitary, cylindrical, erect, brownish green with greyish black trichomes at emergence, copperish green with greyish black pubescent at maturity with an acute apex; peduncle 11 - 16 cm long, 1.5 - 2.0 cm in diameter, erect, tomentose, brown; megasporophylls 56 - 81, 8 - 9 orthostichies with 7 - 9 sporophylls per orthostichy, 4.9 - 5.6 x 2.2 - 2.6 cm, with a prominent distal face, horns straight and robust and 0.63 - 0.81 cm long, 0.95 - 1.35 cm between horns and angle straight between the horns. Seeds 2.43 - 2.71 cm in long, 1.4 - 1.8 cm in diameter, ovoid, sarcotesta whitish pink when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
20036E4799B65E3BA2EF245D63B9AD7C.taxon	distribution	Distribution and habitat. Ceratozamia leptoceras is endemic to the Sierra Madre del Sur in Guerrero State, Mexico (Fig. 19 D), where it occurs on karstic rocks in cloud forest at 1,170 - 1,400 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
20036E4799B65E3BA2EF245D63B9AD7C.taxon	etymology	Etymology. The specific epithet is derived from the Greek words that describe the shape of the horns on the sporophylls: " lepto " for thin or fine and " ceras " in reference to the horns.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
20036E4799B65E3BA2EF245D63B9AD7C.taxon	discussion	Discussion. Ceratozamia leptoceras is distinguished from C. oliversacksii by its linear and membranaceous leaflets. The main differences are in reproductive structures, C. leptoceras has and linear infertile portion of microsporophylls and ovulate strobilus with abundant pubescence at the base of the megasporophylls.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
20036E4799B65E3BA2EF245D63B9AD7C.taxon	materials_examined	Specimens examined. Mexico. Guerrero: Mun. Cochoapa El Grande, 1,170 m, 4 Feb 1984, F. Lorea-Herna ́ ndez 2928 (FCME). Mun. Tlacoachistlahuaca, 1,200 m, 29 May 219, F. Nicolalde-Morejo ́ n et al. 3173 (XAL), 3174 (FCME), 3175 (CIB); 1,200 m, 29 May 2019, L. Marti ́ nez-Domi ́ nguez et al. 1756 (CIB), 1757 (CIB, MEXU), 1758 (XAL), 1759 (CIB, MEXU); 1,400 m, 26 Jun 2019, F. Nicolalde-Morejo ́ n & L. Marti ́ nez-Domi ́ nguez 3255 - 3261 (CIB); 1,400 m, 26 Jun 2019, L. Marti ́ nez-Domi ́ nguez & F. Nicolalde-Morejo ́ n 1860, 1861 (MEXU), 1862 - 1866 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5A1A34A40B3C555F9948FF8DBB39964F.taxon	description	Figs 7 D, 8 D-F, 10 D	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5A1A34A40B3C555F9948FF8DBB39964F.taxon	materials_examined	Type. Mexico. Chiapas: northern slope of Mt. Ovando, 1,000 m, Feb 1939, E. Matuda 2645 ♂ (holotype: MICH! [1002583]; isotypes: CAS! [0001920], MEXU! [acc. # 86830], US! [00620111]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5A1A34A40B3C555F9948FF8DBB39964F.taxon	description	Description. Stem 30 - 50 cm long, 20 - 30 cm in diameter, epigeous, erect and decumbent. Cataphylls 4 - 6.0 x 2.5 - 4.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 3 - 14, 80 - 155 cm long, descending, green at emergence with brown trichomes, glabrous at maturity. Petiole 30 - 79 cm long, terete, linear, green in adult leaves; unarmed to armed with 18 - 36 thin prickles, 0.1 - 0.28 cm long. Rachis 45 - 76 cm long, terete, linear, green with abundant brown trichomes in young leaves, yellow to yellowish green in mature leaves, unarmed to armed with prickles. Leaflets 23 - 53 pairs, opposite to subopposite, insertion in one plane, lanceolate, longitudinally curved abaxially to planar, not basally falcate, papyraceous, flat, green with base yellow and adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 23 - 40 x 0.6 - 1.3 cm, 0.6 - 1.7 cm between leaflets; articulations 0.3 - 0.8 cm wide, yellow. Pollen strobili 9.5 - 16 cm long, 3.5 - 5 cm in diameter, solitary, cylindrical, erect, yellowish green with reddish brown trichomes at emergence, yellowish cream with reddish brown trichomes at maturity; peduncle 8 - 14 cm long, 1.8 - 2.2 cm in diameter, pubescent, reddish brown to brown; microsporophylls 0.78 - 1.4 x 0.77 - 1.1 cm, discoid with a non-recurved distal face and deeply lobate fertile portion, infertile portion 0.37 - 0.56 cm long and orbicular with straight horns 0.16 - 0.35 cm long, 0.50 - 0.81 cm and an acute angle between the horns. Ovulate strobili 8 - 15 cm long, 5 - 8 cm in diameter, solitary, globose, erect, yellowish green with abundant blackish trichomes at emergence, dark green with blackish trichomes at maturity, aristate apex; peduncle 11 - 17 cm long, 1 - 2 cm in diameter, pendulous, with trichomes scarce, blackish, pendulous and erect; megasporophylls 16 - 25, 4 - 5 orthostichies with 4 - 6 sporophylls per orthostichy, 2.5 - 3.4 x 4.5 - 5.3 cm, with a prominent distal face, horns recurved, straight and thin and 0.59 - 1.50 cm long, 0.92 - 1.56 cm between horns with an obtuse angle between the horns. Seeds 2.9 - 4.0 cm long, 1.9 - 4.0 cm in diameter, globose, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5A1A34A40B3C555F9948FF8DBB39964F.taxon	distribution	Distribution and habitat. Ceratozamia matudae is endemic to Chiapas in Mexico (Fig. 20 A), where it occurs on karstic rocks in evergreen tropical forest at 1,000 - 1,500 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5A1A34A40B3C555F9948FF8DBB39964F.taxon	etymology	Etymology. The specific epithet is in honor of professor Eizi Matuda, for his contributions to botany in Mexico.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5A1A34A40B3C555F9948FF8DBB39964F.taxon	discussion	Discussion. Ceratozamia matudae differs from other Ceratozamia species by its yellow rachis and petiole and leaflets with yellow base and articulations. In addition, the length of the peduncle is the same or longer than the fertile portion of ovulate strobili. The ovulate strobili have an aristate apex, and the megasporophylls have long recurved horns.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
5A1A34A40B3C555F9948FF8DBB39964F.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Mun. Acacoyagua, 1,400 m, A. P. Vovides 1069 (XAL); 1,300 m, 9 Jan 1987, D. W. Stevenson et al. 681 (FTG; MO; NY; US); 1,000 m, Feb 1939, E. Matuda 2646 (MEXU); 1,420 m, 24 Jun 2018, F. Nicolalde-Morejon et al. 2832 - 2840 (CIB); 1,420 m, 24 Jun 2018, L. Martinez-Dominguez et al. 1404 - 1409, 1412, 1415, 1416, 1418, 1419 (CIB), 1405, 1407 (CIB, MEXU); 1,480 m, 29 Dec 1993, M. A. Perez-Farrera 27 (CIB, CH, CHIP, MEXU, USCG); 22 Feb 1995, M. A. Perez-Farrera 142 (CHIP, FTG, MEXU). Mun. Escuintla, 8 Feb 2000, O. Farrera S. 1875 (CHIP). Mun. Siltepec, 1,500 m, 6 Nov 1945, E. H. Xolocotzi & A. J. Sharp 367 (MEXU); 1,500 m, Oct-Nov 1940, E. Matuda 4032 (MEXU; NY).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
83C42D63182C5A58A0868E76413D78AB.taxon	description	Figs 7 E, 11 B	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
83C42D63182C5A58A0868E76413D78AB.taxon	materials_examined	Type. Cultivated in Paris " Hort. Bot. Parisiensis ", original material sent by A. Ghiesbrecht from Mexico, 1845, Anon. s. n. (holotype: P! [P 02441737, P 01637464 - P 01637466]). Mexico. Veracruz: Mun. Totutla, Barranca El Coyotito, 1 km road from El Mirador to Conejos, 900 m, 8 Sep 1982, J. Rees & A. P. Vovides 1688 (epitype, designated by Vovides et al. 2016, pg. 427: XAL! [acc. # 17004]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
83C42D63182C5A58A0868E76413D78AB.taxon	description	Description. Stem 20 - 80 cm long, 20 - 45 cm in diameter, epigeous, erect and decumbent. Cataphylls 2 - 6 x 2.5 - 6 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 5 - 55, 100 - 270 cm long, descending, green at emergence, with brown trichomes, glabrous at maturity. Petiole 30 - 93 cm long, terete, linear, dark green in mature leaves; with 8 - 30 thin prickles, 0.06 - 0.20 cm long. Rachis 56 - 154 cm long, terete, linear, green in mature leaves, with prickles and unarmed. Leaflets 25 - 42 pairs, opposite to subopposite, insertion in one plane, lanceolate, in general longitudinally planar, basally falcate, coriaceous, flat, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 29 - 51 x 2.3 - 3.7 cm, 1.8 - 4 cm between leaflets; articulations 0.6 - 1.5 cm wide, green. Pollen strobili 20 - 40 cm long, 4.5 - 7.6 cm in diameter, generally solitary (1 - 2), cylindrical, erect, greenish yellow at emergence, greenish yellow with blackish trichomes at maturity; peduncle 3.5 - 9 cm long, 1.5 - 2.3 cm in diameter, tomentose, reddish brown to light-brown; microsporophylls 1.5 - 2.7 x 0.9 - 2.1 cm, obconic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.47 - 0.68 cm long and orbicular with straight horns 0.29 - 0.40 cm long, 0.51 - 1.05 cm and an acute angle between the horns. Ovulate strobili 23.5 - 40 cm long, 8 - 15 cm in diameter, generally solitary (1 - 2), cylindrical, erect, dark green with blackish trichomes at emergence, dark green with black and gray trichomes at maturity, acuminate apex; peduncle 7.6 - 11.5 cm long, 2.5 - 3 cm in diameter, tomentose, brown to reddish brown, erect or pendulous; megasporophylls 64 - 240, 8 - 16 orthostichies with 8 - 15 sporophylls per orthostichy, 2.0 - 2.7 x 3.2 - 5.3 cm, with a prominent distal face, horns straight and 0.53 - 1.2 cm long, 1 - 2.1 cm between horns and a right angle between the horns. Seeds 2.0 - 3.3 cm long, 1.4 - 2.5 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
83C42D63182C5A58A0868E76413D78AB.taxon	distribution	Distribution and habitat. Ceratozamia mexicana is endemic to Veracruz State, Mexico from the River La Antigua drainage system to the southern end of the Sierra Madre Oriental (Fig. 20 B). It occurs in cloud forest on karstic rocks at 500 - 1,100 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
83C42D63182C5A58A0868E76413D78AB.taxon	etymology	Etymology. The specific epithet is derived from the country of origin of the material used for the description of this species. Ceratozamia mexicana was the first species described in the genus.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
83C42D63182C5A58A0868E76413D78AB.taxon	discussion	Discussion. Ceratozamia mexicana is similar to C. tenuis, C. morettii and C. brevifrons, but it can be easily distinguished from vegetative and reproductive chracters. This species has lanceolate leaflets, whereas C. tenuis and C. morettii have linear and oblong leaflets, respectively. In addition, C. mexicana has insertion in one plane for leaflets and thin prickles on petiole, whereas C. brevifrons has keeled leaflets and petioles armed with robust prickles.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
83C42D63182C5A58A0868E76413D78AB.taxon	materials_examined	Specimens examined. Mexico. Veracruz: Mun. Coatepec, 1,600 m, 26 Jun 1990, P. Zamora C. 2450 (MEXU, XAL). Mun. Comapa, 1,003 m, 18 Feb 2013, F. Nicolalde-Morejon et al. 1732 - 1734 (CIB); 1,003 m, 28 Sep 2014, F. Nicolalde-Morejon et al. 2145 - 2156 (CIB); 970 m, 13 Nov 2015, F. Nicolalde-Morejon et al. 2269 - 2272 (CIB); 1,003 m, 28 Sep 2014, L. Martinez-Dominguez et al. 164 (CIB); 970 m, 13 Nov 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 716 - 723, 725 - 730 (CIB), 724 (CIB, MEXU). Mun. Puente Nacional, 600 m, 13 Mar 1985, G. Castillo-Campos & M. E. Medina 4299 (XAL). Mun. Sochiapa, 1,058 m, 9 Jul 2008, M. Vazquez-Torres 8589 (CIB). Mun. Teocelo, 900 m, 16 Feb 1997, L. H. Bojorquez-Galvan et al. 531 (CIB); 1,070 m, 20 Nov 2015, F. Nicolalde-Morejon et al. 2273 - 2278 (CIB); 1,065 m, 20 Nov 2015, L. Martinez-Dominguez et al. 731 (CIB); 1,070 m, 20 Nov 2015, L. Martinez-Dominguez et al. 732 - 733, 735 - 750 (CIB), 734 (CIB, MEXU); 1,070 m, 11 Mar 2016, L. Martinez-Dominguez et al. 764, 765, 767 - 770 (CIB), 766 (CIB, MEXU); 740 - 800 m, 20 Jul 1995, M. Vazquez-Torres 4865 (CIB). Mun. Tlaltetela, 1,196 m, 5 Jun 2015, F. Nicolalde-Morejon & L. Martinez-Dominguez 2242 (CIB); 1,084 m, 5 Jun 2015, F. Nicolalde-Morejon & L. Martinez-Dominguez 2243 (CIB); 1,196 m, 10 Jun 2015, F. Nicolalde-Morejon & L. Martinez-Dominguez 2246 (CIB); 1,130 m, 21 Jun 2015, F. Nicolalde-Morejon & L. Martinez-Dominguez 2253 - 2257 (CIB); 1,084 m, 5 Jun 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 584 - 586 (CIB); 1,196 m, 10 Jun 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 620 (CIB); 1,130 m, 21 Jun 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 628 - 635 (CIB). Mun. Totutla, 900 m, 23 Feb 1982, A. P. Vovides 730 - 733 (XAL); 900 m, 8 Sep 1982, A. P. Vovides 748 (XAL); 900 m, 9 Oct 1993, Brigada T. Walters s / n (XAL); 875 m, 8 Aug 2015, F. Nicolalde-Morejon et al. 2262 (CIB); 864 m, 27 Jan 2016, F. Nicolalde-Morejon et al. 2279 - 2281 (CIB); F. Vazquez B. 730 (XAL); 1,094 m, 21 Jun 2016, J. M. Ramirez-Amezcua & A. Paizanni Guillen 715 (MEXU); 8 Sep 1982, J. Rees 6344 (IEB); 800 m, 30 Dec 1975, J. Rees & A. P. Vovides 1660, 1672 (XAL); 900 m, 8 Sep 1982, J. Rees & A. P. Vovides 1689 (XAL); 875 m, 8 Aug 2015, L. Martinez-Dominguez et al. 704 - 707 (CIB); 864 m, 27 Jan 2016, L. Martinez-Dominguez et al. 752, 754 - 757 (CIB, MEXU), 758 (CIB). Mun. Xico, 1,195 m, 10 Jul 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 640 - 646 (CIB); 1,159 m, 10 Jul 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 647, 648 (CIB); 900 m, 13 Nov 1981, M. Nee 23035 (NY).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
19E2209773565A98A0C63E75C328118A.taxon	description	Figs 7 F, 8 A-C, 9 D, 11 A	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
19E2209773565A98A0C63E75C328118A.taxon	materials_examined	Type. Mexico. Veracruz: West of Santiago Tuxtla, Cerro el Vigi ́ a, 5 Jul 1983, D. W. Stevenson 542 F (neotype, designated by Stevenson and Sabato 1986, pg. 580: NY! [00001118]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
19E2209773565A98A0C63E75C328118A.taxon	description	Description. Stem 45 - 75 cm long, 16 - 18 cm in diameter, epigeous, erect and decumbent. Cataphylls 3 - 7 x 1.5 - 4 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 12 - 37, 35 - 261 cm long, ascending, light green and glaucous at emergence with brown trichomes, glabrous at maturity. Petiole 31 - 97 cm long, terete, linear, pink at emergence, green in mature leaves; with 8 - 60 robust prickles, 0.22 - 0.59 cm long. Rachis 41 - 164 cm long, terete, linear, pink at emergence, yellowish green in mature leaves, with prickles. Leaflets 12 - 23 pairs, opposite to subopposite, insertion in one plane, oblong, in general longitudinally planar, not basally falcate, papyraceous, flat, green with abaxial and adaxial sides glabrous, acuminate and asymmetrical at the apex, distal end with entire margins, attenuate at base, with conspicuous and green-light veins; median leaflets 16 - 36 x 4.4 - 8.7 cm, 2.9 - 8.5 cm between leaflets; articulations 0.3 - 1.8 cm wide, greenish. Pollen strobili 15 - 30 cm long, 3.1 - 4.2 cm in diameter, usually solitary (1 - 2), cylindrical, erect, greenish yellow with red trichomes at emergence, greenish yellow to cream at maturity; peduncle 3.5 - 5.0 cm long, 1.9 - 2.2 cm in diameter, tomentose, light brown; microsporophylls 1 - 2 x 0.7 - 1.5 cm, obconical with a non-recurved distal face and deeply lobate fertile portion, infertile portion 0.35 - 0.50 cm long and rounded with straight horns 0.30 - 0.42 cm long, 0.52 - 0.70 cm and an acute angle between the horns. Ovulate strobili 22 - 30 cm long, 8.9 - 12 cm in diameter, solitary, cylindrical, erect, green with blackish trichomes, yellowish green with blackish trichomes at maturity, acuminate apex; peduncle 7 - 12 cm long, 2.7 - 3.3 cm in diameter, erect and pendulous, tomentose, light brown; megasporophylls 64 - 110, 8 - 11 orthostichies with 8 - 10 sporophylls per orthostichy, 2.0 - 3.0 x 3.5 - 4.2 cm, with a prominent distal face, horns straight and 0.35 - 0.43 cm long, 1.06 - 1.12 cm between horns with a right angle between the horns. Seeds 2.4 - 3 cm long, 1.3 - 1.8 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
19E2209773565A98A0C63E75C328118A.taxon	distribution	Distribution and habitat. Ceratozamia miqueliana is endemic to southwest Mexico in Chiapas, Tabasco and Veracruz States (Fig. 20 C), where it occurs on shallow soils in evergreen tropical rain forest at 19 - 1,000 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
19E2209773565A98A0C63E75C328118A.taxon	etymology	Etymology. The specific epithet was assigned in honor of Friedrich Anton Wilhelm Miquel.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
19E2209773565A98A0C63E75C328118A.taxon	discussion	Discussion. Ceratozamia miqueliana differs from its congeners by its robust and long prickles, and papyraceous, oblong and glabrous (lustrous) leaflets. In addition, ovulate strobili are yellowish yellowish green with blackish trichomes and have an acuminate apex at maturity. This species is closely related to C. zoquorum, however, that species has descending leaves with coriaceous leaflets.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
19E2209773565A98A0C63E75C328118A.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Mun. Ocozocoautla de Espinosa, 421 m, 23 Oct 1997, R. Jua ́ rez-Galdamez 4 (CHIP, MO, XAL); 800 m, 21 Jan 2001, S. Avendan ̃ o 5223 (MEXU); 765 m, 21 Jan 2001, T. W. Walters 2001 - 2036 - A (MEXU); 2001 - 2036 - C (XAL). Tabasco: Mun. Huimanguillo, 405 m, 10 Aug 2013, F. Nicolalde-Morejo ́ n et al. 1826 - 1846 (CIB); 23 Jun 1998, G. Orti ́ z G. 5291 (MEXU). Veracruz: Mun. A ́ ngel R. Cabada, 231 m, 29 Oct 2013, F. Nicolalde-Morejo ́ n et al. 1868 - 1887 (CIB); 30 Oct 2013, F. Nicolalde-Morejo ́ n et al. 1888 - 1909 (CIB). Mun. Catemaco, 31 Oct 2013, F. Nicolalde-Morejo ́ n et al. 1910 - 1929 (CIB). Mun. Coatzacoalcos, 22 m, 2 Apr 2003, C. H. Ramos 2266 (MEXU); Jun 1960, E. Hernandez X. et al. 162 (CHAPA); 26 Jun 1997, G. Castillo C. & F. Morocini 16267 (XAL); 30 m, 7 Sep 2005, 30 m, L. H. Bojorquez-Galvan 1601 (CIB); 18 Jan 2001, S. Avendan ̃ o R. 5214 (MEXU); 31 m, 18 Jan 2001, T. W. Walters 2001 - 21 - A, B (XAL); T. W. Walters 2001 - 21 - D (MEXU). Mun. Ixhuatlan del Sureste, 45 m, 13 Sep 2011, J. Calonico-Soto & B. Gomez C. 27780 (MEXU). Mun. Las Choapas, 115 m, 26 Nov 2011, A. Rinco ́ n G. 2894 (XAL); 115 m, 30 Jun 2003, E. Lo ́ pez P. 634 (XAL); 100 m, 25 Nov 2004, F. Nicolalde-Morejo ́ n et al. 1434 (XAL). Mun. Mecayapan, 13 Jul 1994, G. Castillo-Campos et al. 12,565 (ENCB, XAL); 9 Mar 1995, G. Castillo-Campos et al. 13470, 13489 (XAL); 5 Aug 1985, J. I. Calzada 11207 (CH, MEXU); 300 m, 16 Jul 1982, M. Nee et al. 25066 (XAL), 25118 (MO, NY, XAL); 950 m, 26 Jan 1992, M. Va ́ zquez- Torres 4123 (CIB); 5 May 1995, M. Va ́ zquez- Torres 5017 (CIB). Mun. Moloacan, 60 m, 19 Dec 1974, J. Rees 1657 (MEXU, XAL), 1658 (XAL), 6347, 6348 (IEB). Mun. Nanchital, 34 m, 12 Aug 2011, J. Rivera H. et al. 4530 (XAL). Mun. Santiago Tuxtla, 900 - 1,000 m, 5 Jul 1983, D. W. Stevenson et al. 541 A-C- F, I, K (NY), G, H, J, N (MEXU, NY); 5 Jul 1983, D. W. Stevenson et al. 542 A-E, H (NY); 650 m, 28 Mar 2014, F. Nicolalde-Morejo ́ n & L. Marti ́ nez-Domi ́ nguez 1988 - 1998 (CIB); 950 m, 24 Jan 1972, J. H. Beaman 5507 (XAL); 650 m, 28 Mar 2014, L. Marti ́ nez-Domi ́ nguez & F. Nicolalde-Morejo ́ n 88 - 106 (CIB), 107 (CIB, MEXU); 800 m, 28 May 1967, L. Scheinvar 673 (MEXU); 29 Aug 1967, M. Sousa 3201 (MEXU); 845 m, 17 Jan 2001, S. Avendan ̃ o R. 5207 (MEXU); 845 m, 17 Jan 2001, T. W. Walters et al. TW 2001 - 2019 (MEXU, XAL). Mun. Pajapan, 830 - 980 m, 15 Jul 1982, M. Nee et al. 25066 (XAL). Mun. Soteapan, 250 m, 13 May 1986, M. Va ́ zquez-Torres 3360 (CIB, XAL). Mun. Tatahuicapan de Jua ́ rez, 849 m, C. I. Carvajal & M. J. Fragoso 797 (CIB); 15 Mar 2008, L H. Bojorquez-Galvan et al. 1939, 1946 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
E90B352E7ECE57BA841E097AA650C2F5.taxon	description	Fig. 7 G	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
E90B352E7ECE57BA841E097AA650C2F5.taxon	materials_examined	Type. Mexico. Chiapas: Mun. Villa Flores, Ejido La Sombra de la Selva, 880 m, 20 Sep 1997, J. De La Cruz Rodriguez 66 (lectotype, designated here: MEXU! [MEXU 00934924, MEXU 00934907, MEXU 00934905]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
E90B352E7ECE57BA841E097AA650C2F5.taxon	description	Description. Stem 20 - 60 cm long, 20 - 30 cm in diameter, epigeous, erect and decumbent. Cataphylls 3 - 8.0 x 1.5 - 4.0 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 2 - 50 (150), 46 - 242.5 cm long, ascending, brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole 16 - 98 cm long, terete, linear, brown in mature leaves; with 11 - 46 thin prickles, 0.30 - 0.40 cm long. Rachis 26 - 144.5 cm long, terete, linear, brown to greenish brown in mature leaves, lower 2 / 3 rds with prickles. Leaflets 28 - 94 pairs, opposite to subopposite, insertion in one plane, linear to lanceolate, in general longitudinally planar, not basally falcate, papyraceous, lightly involute, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 10 - 40 x 0.4 - 0.9 cm, 0.2 - 1.6 cm between leaflets; articulations 0.12 - 1.20 cm wide, brown. Pollen strobili 16 - 55 cm long, 4.0 - 7.7 cm in diameter, solitary, cylindrical, erect, yellowish green with brown trichomes at emergence, yellowish cream with blackish trichomes at maturity; peduncle 5 - 11 cm long, 0.8 - 1.4 cm in diameter, pubescent, reddish brown to brown; microsporophylls 1.23 - 1.80 x 0.62 - 0.86 cm, elliptic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.62 - 0.81 cm long and linear with straight horns 0.17 - 0.39 cm long, 0.56 - 0.95 cm and an obtuse to acute angle between the horns. Ovulate strobili 13.5 - 35 cm long, 7 - 12.6 cm in diameter, solitary, cylindrical, erect, yellowish green with abundant blackish trichomes at emergence, glaucous green with reddish brown to blackish trichomes at maturity, acuminate apex; peduncle with trichomes scarce, brownish, 4.5 - 12 cm long, 1.0 - 2.2 cm in diameter, erect; megasporophylls 42 - 72, 6 - 8 orthostichies with 7 - 10 sporophylls per orthostichy, 2.0 - 3.0 x 2.8 - 6.1 cm, with a prominent distal face, horns straight and 0.29 - 1.47 cm long, 1.55 - 2.09 cm between horns with an obtuse angle between the horns. Seeds 1.9 - 2.9 cm long, 1.2 - 2.5 cm in diameter, spherical, sarcotesta whitish pink when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
E90B352E7ECE57BA841E097AA650C2F5.taxon	distribution	Distribution and habitat. Ceratozamia mirandae is widely distributed in Chiapas State, Mexico from " Tres Picos " hill and Sierra Morena to a mountain range in Mun. Jiquipilas (Fig. 20 D); it occurs on karstic rocks with abundant organic material in oak forest and transitional oak forest between pine oak forest and cloud forest between 850 and 1,500 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
E90B352E7ECE57BA841E097AA650C2F5.taxon	etymology	Etymology. This species was named in honor of Dr. Faustino Miranda for his untiring contributions to the flora of Chiapas (Vovides et al. 2001).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
E90B352E7ECE57BA841E097AA650C2F5.taxon	discussion	Discussion. Ceratozamia mirandae is similar to C. alvarezii. However, in C. mirandae, the leaves and ovulate strobilus are larger at the population level than in C. alvarezii, which is a more diminutive species overall. In the original publication of C. mirandae, there were no isotypes cited, only the holotype in CHIP. Because no holotype of C. mirandae has been found in CHIP, despite intensive searches, we designate the uncited isotype found in MEXU as the lectotype.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
E90B352E7ECE57BA841E097AA650C2F5.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Mun. Jiquipilas, 1,170 m, 19 Jun 2018, F. Nicolalde-Morejon et al. 2749 - 2759 (CIB); 1,015 m, 8 Mar 1995, J. J. Castillo-Hernandez 548 (CHIP); 1,170 m, 19 Jun 2018, L. Martinez-Dominguez et al. 1312 - 1316, 1318, 1320, 1321 (CIB), 1317 (CIB, MEXU), 1319 (CIB, MEXU); 1,270 m, 25 May 1995, M. A. Perez-Farrera 465 (CHIP). Mun. Villa Corzo, 1,320 m, 12 Jul 2004, A. Reyes-Garcia et al. 7134 (MEXU); 1,500 m, 9 Feb 1972, D. E. Breedlove 23999 (MEXU, MO); 1,170 m, 16 Mar 1989, U. Bachem & R. Rojas C. 579 (CHIP, MEXU, SLPM). Mun. Villaflores, 960 m, 6 Apr 1995, A. P. Vovides & M. A. Perez-Farrera 1261 (XAL); 1,157 m, 7 Jul 2004, D. Alvarez 9809 (MEXU); 1,277 m, 25 Jun 2018, F. Nicolalde-Morejon et al. 2854 - 2863 (CIB); 1,195 m, 20 Jun 2019, F. Nicolalde-Morejon & L. Martinez-Dominguez 3208 - 3213 (CIB); 1,350 m, 21 Jun 2019, F. Nicolalde-Morejon & L. Martinez-Dominguez 3215 - 3224 (CIB); 1,015 m, 5 Aug 1994, J. J. Castillo-Hernandez 230 (CIB, MEXU, USCG); 1,250 m, 5 Apr 1995, J. J. Castillo-Hernandez 595 (CHIP); 1,140 m, 6 Aug 2002, L. Alvarado C. et al. 368 (MEXU); 1,195 m, 20 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1808 - 1813 (CIB); 1,350 m, 21 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1814 - 1825 (CIB); 1,277 m, 25 Jun 2018, L. Martinez-Dominguez et al. 1428, 1429, 1431 - 1438 (CIB), 1430 (CIB, MEXU); 940 m, 22 Dec 1993, M. A. Perez-Farrera s / n (XAL), 26 (CHIP, XAL); 910 m, 28 Apr 1994, M. A. Perez-Farrera 37 (CHIP, XAL); 1,460 m, 16 Sep 1994, M. A. Perez-Farrera 126 (CHIP); 950 m, 12 Apr 1996, M. A. Perez-Farrera 1480 (CHIP).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AF657135F53A5B7BBCCD310DF063E586.taxon	description	Fig. 7 H	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AF657135F53A5B7BBCCD310DF063E586.taxon	materials_examined	Type. Mexico. Oaxaca: Cercania a Juquila Mixe, 1,737 m, 21 May 1997. J. Chemnick & T. Gregory 49 (lectotype, designated by Nicolalde-Morejon and Avendano 2011, pg. 1033: XAL! [XAL 0065870]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AF657135F53A5B7BBCCD310DF063E586.taxon	description	Description. Stem 30 - 100 cm long, 18 - 25 cm in diameter, epigeous, decumbent. Cataphylls 1.8 - 4.6 x 1.9 - 4.3 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 3 - 17 (30), 85 - 246 cm long, descending, light green and glaucous at emergence with light brown trichomes, glabrous at maturity. Petiole 46 - 132 cm long, terete, linear, green in mature leaves; with 30 - 50 thin prickles, 0.20 - 0.38 cm long. Rachis 58 - 118 cm long, terete, linear, green to greenish in mature leaves, with scarce prickles. Leaflets 19 - 35 pairs, opposite to subopposite insertion in one plane, lanceolate, abaxially curved, rarely planar, not basally falcate, papyraceous, flat, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 24 - 39 x 1.9 - 2.9 cm, 1.7 - 3.0 cm between leaflets; articulations 0.6 - 1.0 cm wide, green. Pollen strobili 22 - 30 cm long, 4.5 - 7 cm in diameter, generally solitary (1 - 2), cylindrical, erect, greenish at emergence with reddish trichomes, greenish yellow with reddish brown trichomes at maturity; peduncle 10 - 15 cm long, 1.2 - 2.0 cm in diameter, glabrous or with trichomes scarce reddish brown to brown; microsporophylls 1.4 - 2.4 x 0.7 - 1.3 cm, obconic with a non-recurved distal face and lobate fertile portion, infertile portion 0.39 - 0.48 cm long and linear with straight horns 0.30 - 0.50 cm long, 0.50 - 0.93 cm and a right angle between the horns. Ovulate strobili 23 - 32 cm long, 12 - 16 cm in diameter, solitary, cylindrical, erect, yellowish green with abundant blackish trichomes at emergence, green with brown to blackish trichomes at maturity, apiculate apex; peduncle 12 - 23.5 cm long, 1.3 - 2.1 cm in diameter, pendulous, glabrous or with scarce reddish brown trichomes; megasporophylls 60 - 112, 6 - 9 orthostichies with 10 - 14 sporophylls per orthostichy, 2.4 - 2.8 x 4.0 - 5.1 cm, with a truncate distal face, horns straight and 0.90 - 1.1 cm long, 0.92 - 1.2 cm between horns and a right angle between the horns. Seeds 2.2 - 3.0 cm long, 1.6 - 2.0 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AF657135F53A5B7BBCCD310DF063E586.taxon	distribution	Distribution and habitat. Ceratozamia mixeorum is endemic to the southern portion of Sierra Norte of Oaxaca State (Mexico), along the highlands in the Mixe area (Fig. 21 A), where it occurs in karstic rocks of cloud and oak forests between 1,170 to 2,150 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AF657135F53A5B7BBCCD310DF063E586.taxon	etymology	Etymology. The specific epithet was named in honor of the Mixe people.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AF657135F53A5B7BBCCD310DF063E586.taxon	discussion	Discussion. Ceratozamia mixeorum is similar to C. whitelockiana, but has prickles on the rachis of the leaf and a densely prickly petiole (up to 50 prickles), ovulate strobili with long and pendulous pedunclse up to 23.5 cm long. The fertile portion of ovulate strobilus has 6 - 9 orthostichies and 10 - 14 sporophylls per orthostichy; in contrast to 4 - 5 orthostichies and 5 - 8 sporophylls per orthostichy in C. whitelockiana.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AF657135F53A5B7BBCCD310DF063E586.taxon	materials_examined	Specimens examined. Mexico. Oaxaca: Mun. Guevea de Humboldt, 1,300 m, 30 Mar 1991, A. Campos 3614 (MEXU). Mun. San Juan Juquila Mixes, 1,605 m, 7 Apr 1995, E. Torres B. 563 (SERO, XAL); 1,595 m, 25 Sep 2020, F. Nicolalde-Morejon et al. 3368 (CIB); 1,681 m, 25 Sep 2020, F. Nicolalde-Morejon et al. 3369 - 3371 (CIB); 1,595 m, 25 Sep 2020, L. Martinez-Dominguez et al. 2001, 2002 (CIB); 1,681 m, 25 Sep 2020, L. Martinez-Dominguez et al. 2003 (CIB, MEXU), 2004 - 2006 (CIB). Mun. San Pedro Ocotepec, 8 Apr 1995, 1,600 m, E. Torres B. et al. 607 (XAL). Mun. Santa Maria Guienagati, 1,200 m, 25 Jan 1996, M. Ceron C. et al. 249 (MEXU, SERO, XAL). Mun. Santa Maria Tepantlali, 1,728 m, 25 Sep 2020, F. Nicolalde-Morejon et al. 3362 - 3367 (CIB); 2,147 m, 15 May 2011, G. Juarez-Garcia 3871 (MEXU); 1,728 m, 25 Sep 2020, L. Martinez-Dominguez et al. 1994 - 1995 (CIB, MEXU), 1996 - 2000 (CIB). Mun. Totontepec Villa de Morelos, 1,300 m, 7 Dec 1989, E. Vargas-Ruiz 476 (MEXU); 1,900 m, 21 Apr 1986, J. Rivera-Reyes & Gary J. Martin 246 (MEXU); 1,700 m, 15 Jul 1987, E. Velasco-Lopez & G. J. Martin 60 (MEXU); 1,900 m, 6 Jan 1988, J. Rivera-Reyes & G. J. Martin 920 (MEXU); 2,000 m, 16 Sep 1987, W. Ruiz S. 46 (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
122290CE5CE35A3DB71C802B069282A4.taxon	description	Figs 7 I-J, 10 A, C	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
122290CE5CE35A3DB71C802B069282A4.taxon	materials_examined	Type. Mexico. Veracruz: Mun. Landero y Coss, El Raicero, 4 km NE from Landero y Coss, 1,500 m, 7 Jan 1992, M. Va ́ zquez-Torres & H. Barney 4097 (holotype: CIB! [acc. # 22297 UV]; isotypes: CIB! [acc. # 24578 UV], MEXU [n. v.], XAL! [XAL 0001061, XAL 0001064])	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
122290CE5CE35A3DB71C802B069282A4.taxon	description	Description. Stem 20 - 23 (50) cm long, 20 - 35 cm in diameter, epigeous, erect and decumbent. Cataphylls 4 - 6.5 x 3 - 3.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 3 - 30, 82 - 200 cm long, descending, light green at emergence, with brown trichomes, glabrous at maturity. Petiole 30 - 90 cm long, terete, linear, green in mature leaves; with 8 - 60 thin prickles, 0.01 - 0.20 cm long. Rachis 50 - 116 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 10 - 23 pairs, opposite to subopposite, oblong, longitudinally curved abaxially to planar, not basally falcate, coriaceous, green with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acuminate and asymmetrical at the apex, attenuate at base, and with conspicuous and green-light veins; median leaflets 17.5 - 41 x 2.6 - 4.3 cm wide, 2.1 - 6 cm between leaflets; articulations 0.5 - 1.6 cm wide, green and yellow. Pollen strobili 12 - 22 (35.5) cm long, 3.8 - 5.2 cm in diameter, solitary, cylindrical, erect, brownish yellow at emergence, greenish yellow with blackish pubescent at maturity; peduncle 3 - 7 (12.5) cm long, 1.1 - 1.6 cm in diameter, tomentose, reddish brown to brown; microsporophylls 1.0 - 2.0 x 0.9 - 1.15 cm, obconic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.37 - 0.75 cm long and orbicular with straight horns 0.15 - 0.53 cm long, 0.47 - 0.92 cm and an acute angle between the horns. Ovulate strobili 14 - 20 cm long, 8 - 9.9 cm in diameter, solitary, globose, erect, yellowish green with brown trichomes at emergence, green with blackish trichomes at maturity and with apiculate apex; peduncle 4 - 7 cm long, 1.2 - 1.8 cm in diameter, erect, tomentose, brown; megasporophylls 35 - 81 (108), 7 - 10 orthostichies with 5 - 9 sporophylls per orthostichy, 1.7 - 2.6 x 3.0 - 4.5 cm, with a truncate distal face, horns straight and 0.32 - 0.69 cm long, 1.05 - 1.6 cm between horns and a right angle between the horns. Seeds 1.5 - 2.3 cm, 1 - 1.6 cm in diameter, ovate and globose, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
122290CE5CE35A3DB71C802B069282A4.taxon	distribution	Distribution and habitat. Ceratozamia morettii is endemic to the Sierra de Chiconquiaco in Veracruz State, Mexico (Fig. 21 B), here it occurs in cloud forest on karstic rocks and cliffs between 1,150 - 1,850 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
122290CE5CE35A3DB71C802B069282A4.taxon	etymology	Etymology. The specific epithet honors the Italian botantist Aldo Moretti from the Orto Botanico and Istituto di Biologia Vegetale at the University of Naples Federico II, in recognition of his scientific contributions to cycad biology (Vazquez-Torres et al. 1998).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
122290CE5CE35A3DB71C802B069282A4.taxon	discussion	Discussion. In leaf morphology, Ceratozamia morettii is morphologically similar to C. delucana; however, there are differences in the total size of the plant and in its reproductive structures. The microsporophyll horns of C. morettii form an acute angle, whereas those of C. delucana form a right angle. The ovulate strobilus in C. morettii is green with blackish trichomes at maturity and an apiculate apex. In contrast, in C. delucana, the ovulate strobilus is green and glabrous at maturity with an acute apex.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
122290CE5CE35A3DB71C802B069282A4.taxon	materials_examined	Specimens examined. Mexico. Veracruz: Mun. Chiconquiaco, 1,700 m, 5 Sep 1981, A. P. Vovides 687 (XAL); 1,850 m, 18 Dec 1981, A. P. Vovides 704 (XAL); Sep 1829, C. J. W. Schiede s / n (XAL); 1,598 m, 17 Sep 2020, F. Nicolalde-Morejon et al. 3318 - 3322 (CIB); 26 Nov 1974, J. Rees 6336 (IEB); 1,700 m, 5 Jun 2018, L. Lagunes G. 83, 84, 85, 86 (CIB); 1,600 m, 26 Apr 2018, L. Martinez-Dominguez et al. 1280 - 1290 (CIB); 1,700 m, 3 Dec 2018, L. Martinez-Dominguez et al. 1567, 1568 (CIB); 1,500 m, 15 Mar 2019, L. Martinez-Dominguez et al. 1660, 1661 (CIB); 1,550 m, 26 Mar 2019, L. Martinez-Dominguez et al. 1680, 1681 (CIB); 1,598 m, 17 Sep 2020, L. Martinez-Dominguez et al. 1931 - 1934, 1936 (CIB), 1935 (CIB, MEXU). Mun. Landero y Coss, 1,750 m, 1 Jan 1976, J. Rees & A. P. Vovides 1662 - 1664 (XAL); 1,750 m, 23 Sep 1976, J. Rees & A. P. Vovides 1676 (XAL); 1,830 m, 3 Mar 2011, L. H. Bojorquez G. et al. 23101 - 2316 (CIB); 1,472 m, 29 Jan 2015, L. Martinez-Dominguez et al. 185 - 214 (CIB); 1,765 M, 22 Apr 2008, M. Vazquez-Torres et al. 8349 (CIB); 3 Mar 2001, S. Avendano 5378 (MEXU); 1,520 m, 10 Jan 2001, T. W. Walters 2001 - 01 - E (XAL). Mun. Tenochtitlan, 1,500 m, 10 Apr 2002, A. Rincon G. et al. 2996, 2997 (XAL); 1,500 m, 12 Apr 2002, A. Rincon G. et al. 2998 (XAL). Mun. Yecuatla, 1,400 m, 26 May 1981, C. Gutierrez B. & A. Montoya L. 134 (MO, XAL); 1,211 m, 28 Sep 2014, F. Nicolalde-Morejon & L. Martinez-Dominguez 2087 - 2106 (CIB); 1, 200 m, 26 Sep 1976, J. Rees 1677 (XAL); 1,211 m, 9 Aug 2014, L. Martinez-Dominguez et al. 161 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9CBD0D5C557252D3953F80AA0BABEA34.taxon	description	Fig. 27 A	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9CBD0D5C557252D3953F80AA0BABEA34.taxon	materials_examined	Type. Mexico. Chiapas: Rancho Fenia, Mar-Apr 1925, C. Purpus 6 ♀ (holotype: NY! [00001116]; isotypes: F! [acc. # 1530231], MO!, US! [00620294]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9CBD0D5C557252D3953F80AA0BABEA34.taxon	description	Description. Stem 30 - 60 cm long, 20 - 40 cm in diameter, epigeous, erect and decumbent. Cataphylls 4.5 - 6 x 2.5 - 6.4 cm wide at the base, persistent, long triangular, reddish brown, densely brown tomentose at emergence, tomentose at maturity, apex acuminate. Leaves 5 - 48, 50 - 126 cm long, ascending, copperish brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole 10 - 49 cm long, terete, twisted, copperish with abundant greyish trichomes in young leaves, copperish and glabrous in mature leaves; with 30 - 60 robust prickles, 0.11 - 0.31 cm long. Rachis 40 - 88 cm long, terete, twisted, copperish with abundant greyish trichomes in young leaves, copperish and glabrous in mature leaves, with prickles. Leaflets 33 - 91 pairs, opposite to subopposite, insertion irregular to in one plane, linear, in general longitudinally planar, not basally falcate to basally falcate, coriaceous, strongly involute to caniculate, green with yellowish green base and with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acute and symmetric at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 14 - 48 x 0.21 - 0.46 cm, 0.2 - 1.3 cm between leaflets; articulations 0.08 - 0.66 cm wide, yellowish. Pollen strobili 13.2 - 25 cm long, 3.1 - 8 cm in diameter, solitary, cylindrical, erect, brown with reddish brown trichomes at emergence, yellowish green to cream at maturity; peduncle 2.0 - 5.5 cm long, 1.3 - 2.3 cm in diameter, tomentose, brown; microsporophylls 0.92 - 1.16 x 0.86 - 0.1.1 cm, discoid with a non-recurved distal face and a lobate fertile portion, infertile portion 0.36 - 0.45 cm long and orbicular with straight horns 0.07 - 0.13 cm long, 0.35 - 0.56 cm and an acute angle between the horns. Ovulate strobili 21 - 35 cm long, 8.5 - 10.5 cm in diameter, solitary, cylindrical, erect, green with abundant blackish trichomes at emergence, grayish green with abundant blackish trichomes at maturity, acuminate apex; peduncle 4.8 - 9 cm long, 1.8 - 2.2 cm in diameter, erect, tomentose, brown; megasporophylls 42 - 63, 6 - 7 orthostichies with 7 - 9 sporophylls per orthostichy, 2.6 - 3.0 x 3.6 - 4.0 cm, with a truncated distal face, horns straight and 0.37 - 0.58 cm long, 0.37 - 0.42 cm between horns with a right angle between the horns. Seeds 2.0 - 3.5 cm long, 1.1 - 1.9 cm in diameter ovate, sarcotesta whitish pink when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9CBD0D5C557252D3953F80AA0BABEA34.taxon	distribution	Distribution and habitat. Ceratozamia norstogii is distributed in mountainous regions in Chiapas and Oaxaca States, Mexico (Fig. 21 C), where it occurs on karstic rocks in pine-oak forest and the transition between pine forest and oak forest between 800 - 1,650 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9CBD0D5C557252D3953F80AA0BABEA34.taxon	etymology	Etymology. This species was named in honor of Knut Norstog, for his extensive and significant contributions to cycad biology (Stevenson 1982).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9CBD0D5C557252D3953F80AA0BABEA34.taxon	discussion	Discussion. Ceratozamia norstogii is easily diagnosable by its petiole and raquis twisted. The linear leaflets are coriaceous, strongly involute and appear caniculate and have an acute apex.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9CBD0D5C557252D3953F80AA0BABEA34.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Mun. Cintalapa, 1,100 m, 19 Mar 1993, A. P. Vovides et al. 1230, 1231, 1233, 1235 (XAL); 1,600 m, 3 Nov 1971, D. E. Breedlove 21813 (MEXU, MO); 1,600 m, 21 Apr 1972, D. E. Breedlove 24709 (MO); 1,600 m, 12 Oct 1979, D. E. Breedlove 44431 (MEXU); 800 m, 4 May 1988, E. Palacios E. 375 (CHIP, IBUG); 1,240 m, 6 Jun 1993, E. Palacios E. 2155 (CHIP); 1,280 m, 22 May 2001, J. M. Lazaro-Zermeno 251 (CHIP); 1,038 m, 20 Jun 2018, F. Nicolalde-Morejon et al. 2762 - 2770 (CIB); 1,325 m, 20 Jun 2018, F. Nicolalde-Morejon et al. 2771 - 2780 (CIB); 1,038 m, 20 Jun 2018, L. Martinez-Dominguez et al. 1326 - 1334 (CIB), 1335 (CIB, MEXU); 1,325 m, 20 Jun 2018, L. Martinez-Dominguez et al. 1337 - 1349 (CIB); 1,100 m, 5 Oct 1995, M. A. Perez-Farrera 775 (CH, CIB, XAL); 5 Dec 1996, M. A. Perez-Farrera 1483 (HEM). Oaxaca: Mun. San Miguel Chimalapa, 1,120 m, 1 Apr 1996, S. Salas-Morales et al. 1173 (SERO, XAL). Mun. Santo Domingo Zanatepec, 800 m, 22 Jun 2018, F. Nicolalde-Morejon et al. 2819 - 2828 (CIB); 800 m, 22 Jun 2018, L. Martinez-Dominguez et al. 1380, 1381, 1383, 1386, 1389 - 1391, 1394, 1396, 1399 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2644CF65A3C5560F907C3AA4FF9B52EF.taxon	description	Fig. 27 B	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2644CF65A3C5560F907C3AA4FF9B52EF.taxon	materials_examined	Type. Mexico. Oaxaca: Mun. Candelaria Loxicha, Cerro Perico, 1,616 m, 10 Jun 2021, L. Marti ́ nez-Domi ́ nguez et al. 2261 ♀ (holotype: CIB! [acc. # 23411 UV]; isotypes: K!, MEXU!, NY!).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2644CF65A3C5560F907C3AA4FF9B52EF.taxon	description	Description. Stem 30 - 80 cm long, 10 - 40 cm in diameter, epigeous, decumbent. Cataphylls 6.5 - 7.5 x 1.6 - 2.0 cm wide at the base, persistent, triangular, reddish brown, brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 7 - 36 (50), 124 - 258 cm long, descending, green at emergence with brown trichomes, glabrous at maturity. Petiole 45 - 110 cm long, terete, linear, green at emergence, dark green in mature leaves; with 40 - 65 thin prickles, 0.50 - 0.70 cm long. Rachis 85 - 182 cm long, terete, linear, dark green in mature leaves with prickles. Leaflets 24 - 47 pairs, opposite to subopposite, insertion in one plane, linear, longitudinally curved abaxially to planar, not basally falcate, papyraceous, flat, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical (rarely asymmetrical) at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 25 - 40 x 2.3 - 3.6 cm, 2.0 - 3.3 cm between leaflets; articulations 0.5 - 1.1 cm wide, green. Pollen strobili 20 - 35 cm long, 3.0 - 5.7 cm in diameter, solitary, cylindrical, erect, greenish with reddish trichomes at emergence, greenish yellow with reddish brown trichomes at maturity; peduncle 10 - 15 cm long, 1.2 - 1.8 cm in diameter, pubescent, reddish brown; microsporophylls 2.5 - 2.75 x 0.99 - 1.28 cm, obconic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.60 - 0.80 cm long and a rounded with straight horns 0.08 - 0.18 cm long, 0.50 - 0.70 cm and an acute angle between the horns. Ovulate strobili 33 - 36 cm long, 11.5 - 14 cm in diameter, solitary, cylindrical, erect, greenish yellow with few reddish trichomes at emergence, green with brown to reddish brown trichomes or glabrous at maturity, acute apex; peduncle 8.0 - 10.0 cm long, 1.4 - 1.6 cm in diameter, erect, pubescent, light brown; megasporophylls 80 - 210, 9 - 10 orthostichies with 11 - 13 sporophylls per orthostichy, 2.3 - 2.9 x 3.9 - 5.0 cm, with a truncate distal face, horns straight and 0.31 - 0.53 cm long, 1.33 - 1.40 cm between horns with a right angle between the horns. Seeds 2.45 - 2.65 cm long, 1.4 - 1.6 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2644CF65A3C5560F907C3AA4FF9B52EF.taxon	distribution	Distribution and habitat. Ceratozamia oliversacksii occurs along the Eastern Sierra Madre del Sur in the Oaxacan Highlands, Mexico (Fig. 21 D), it is found on karstic rocks with abundant organic matter in cloud forest and the transition zone between it and pine forest between 1,040 to 1,850 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2644CF65A3C5560F907C3AA4FF9B52EF.taxon	etymology	Etymology. The specific epithet honors Oliver Sacks, who loved cycads and was a distinguished American neurologist and historian of science. Sacks published Island of the Color Blind and Cycad Island (Sacks 1997) and Oaxaca Journal (Sacks 2002) (Martinez-Dominguez et al. 2022 b).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2644CF65A3C5560F907C3AA4FF9B52EF.taxon	discussion	Discussion. Ceratozamia oliversacksii is similar to C. robusta and C. leptoceras but is characterized by green leaves at emergence and the combination of linear and papyraceous leaflets. The ovulate strobili have 11 to 13 sporophylls per orthostichies and an acute apex, acute horns of megasporophylls up to 0.53 cm long, and microsporophylls with a rounded distal face and straight horns.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
2644CF65A3C5560F907C3AA4FF9B52EF.taxon	materials_examined	Specimens examined. Mexico. Oaxaca: Mun. Candelaria Loxicha, 1,380 m, 3 May 2005, A. Luna-Jose & B. Rendon-Aguilar 1472, 1473 (XAL); 1,630 m, 23 Jun 2019, F. Nicolalde-Morejon & L. Martinez-Dominguez 3231 - 3243 (CIB); 1,630 m, 23 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1832 - 1839 (CIB), 1840 (CIB, MEXU); 1,418 m, 20 April 2021, L. Martinez-Dominguez et al. 2160, 2161 (CIB); 1,616 m, 10 Jun 2021, M. Rios-Mendez et al. 105 (CIB). Mun. San Agustin Loxicha, 1,760 m, 12 Oct 2003, A. Luna-Jose & B. Rendon-Aguilar 5 18 (XAL); 1,400 m, 23 Jun 2019, F. Nicolalde-Morejon & L. Martinez-Dominguez 3229, 3230 (CIB); 1,400 m, 23 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1829 - 1831 (CIB). Mun. Santa Catarina Juquila, 1,850 m, 13 Apr 1965, J. Rzedowski 1 9557 (ENCB); 1,450 - 1,700 m, 11 Feb 1965, R. McVaugh 22346 (ENCB; MICH). Mun. San Miguel del Puerto, 1,060 m, 25 Jun 2019, F. Nicolalde-Morejon & L. Martinez-Dominguez 3244 - 3249 (CIB); 1,430 m, 17 Apr 2000, J. Rivera H. et al. 2378 (FCME, SERO); 1,060 m, 25 Jun 2019, L. Martinez-Dominguez & F. Nicolalde-Morejon 1847 - 1849, 1851, 1852 (CIB), 1850 (CIB, MEXU); 1,040 m, 1 Apr 2003, L. Schibli et al. 152 (SERO).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
FEC37F523E7B59B2818B8BA674999F43.taxon	description	Figs 5, 6	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
FEC37F523E7B59B2818B8BA674999F43.taxon	diagnosis	Diagnosis. Similar to Ceratozamia robusta and C. subroseophylla, but distinguished by characters of reproductive structures. It has megasporophylls with abundant purple to wine-colored trichomes, ovulate strobili with an acuminate apex and up to 12 orthostichies with 31 sporophylls per orthostichy and pollen strobili with a long infertile portion from 0.65 to 0.80 cm long.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
FEC37F523E7B59B2818B8BA674999F43.taxon	materials_examined	Type. BELIZE. Toledo: Southwestern Maya Mountains, Columbia River Forest Reserve, Union Camp. 6 Apr 1992, B. K. Holst 4105 (holotype: NY! [01340569]; isotype: MO! [acc. # 04661737, 04661738]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
FEC37F523E7B59B2818B8BA674999F43.taxon	description	Description. Stem 30 - 200 cm long, 20 - 40 cm in diameter, epigeous, generally decumbent. Cataphylls 5.5 - 8 x 2 - 3 cm wide at the base, persistent, triangular, reddish brown, brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 12 - 50, 90 - 300 cm long, ascending, brown at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 75 - 100 cm long, terete, linear, greenish brown or dark brown at emergence, green in mature leaves; with 20 - 40 robust prickles, 0.20 - 0.45 cm long. Rachis 90 - 200 cm long, terete, linear, greenish brown or dark brown at emergence, green in mature leaves, with prickles. Leaflets 26 - 51 pairs, opposite to subopposite, insertion in one plane, lanceolate, longitudinally curved abaxially to planar, not basally falcate, papyraceous, flat, dark green, adaxial side glabrous, abaxial side glabrous, distal end with entire margins, acuminate and symmetric at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 30 - 45 x 2.5 - 4.0 cm, 1.7 - 5 cm between leaflets; articulations 0.9 - 1.6 cm wide, brown in young leaves and green in mature leaves. Pollen strobili 60 - 80 cm long, 7 - 9 cm in diameter, solitary, cylindrical, erect, green with wine trichomes at emergence, greenish yellow at maturity with wine to purple trichomes; peduncle 7 - 11 cm long, 2.5 - 3.5 cm in diameter, tomentose, brown; microsporophylls 2.0 - 3.3 x 1.1 - 1.85 cm, obconic with a non-recurved distal face and deeply lobate fertile portion, infertile portion 0.65 - 0.80 cm long and rounded with straight horns 0.20 - 0.29 cm long, 0.55 - 0.75 cm and a right angle between the horns. Ovulate strobili 30 - 50 cm long, 9 - 14 cm in diameter, solitary, cylindrical, erect, green with abundant wine to dark purple trichomes at emergence, green with abundant dark purple trichomes at maturity, acuminate apex; peduncle 12 - 20 cm long, 2.5 - 4.5 cm in diameter, erect, tomentose, reddish brown; megasporophylls 108 - 280, 9 - 12 orthostichies with 12 - 31 sporophylls per orthostichy, 1.8 - 2.3 x 2.3 - 4.1 cm, with a prominent distal face, horns straight and 0.36 - 0.50 cm long, 0.90 - 1.35 cm between horns with a right angle between the horns. Seeds 3.0 - 4.5 cm long, 2.0 - 3.0 cm in diameter, ovate, sarcotesta whitish pink when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
FEC37F523E7B59B2818B8BA674999F43.taxon	distribution	Distribution and habitat. Ceratozamia osbornei is endemic to Belize (Fig. 22 A), where it occurs in evergreen tropical forest on karstic rocks with abundant organic matter between 200 and 750 m elevation.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
FEC37F523E7B59B2818B8BA674999F43.taxon	etymology	Etymology. The specific epithet honors Roy Osborne in recognition of his remarkable contributions to the knowledge of the cycad diversity in the world. In particular, he has actively participated in the project " The World List of Cycads " since its inception (https: // www. cycadlist. org).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
41F0AD37EA8F5A45B4270E4722B73AC4.taxon	description	Fig. 27 C	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
41F0AD37EA8F5A45B4270E4722B73AC4.taxon	materials_examined	Type. Mexico. Chiapas: NW of Tuxtla Gutierrez on road through San Fernando 23.5 km from route 190, 1,000 m, 9 Jul 1983, D. W. Stevenson 549 A (neotype, designated by Stevenson and Sabato 1986, pg. 581: NY! [00001119]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
41F0AD37EA8F5A45B4270E4722B73AC4.taxon	description	Description. Stem 30 - 100 cm long, 20 - 40 cm in diameter, epigeous, erect and decumbent. Cataphylls 5.0 - 9.0 x 3.5 - 7.5 cm wide at the base, persistent, long triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 10 - 50, 138 - 266 cm long, ascending, dark brown at emergence, green and glabrous at maturity. Petiole 56 - 85 cm long, terete, linear, brown in young leaves, green in mature leaves; with 27 - 55 robust prickles, 0.48 - 0.70 cm long. Rachis 60 - 190 cm long, terete, linear, brown in young leaves, green in mature leaves, with prickles. Leaflets 20 - 45 pairs, opposite to subopposite, insertion in one plane, lanceolate, in general longitudinally planar, not basally falcate, papyraceous, flat, green with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acuminate and asymmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 28 - 42 x 2.5 - 4.0 cm, 3.0 - 4.5 cm between leaflets; articulations 0.2 - 0.8 cm wide, green. Pollen strobili 50 - 70 cm long, 7.0 - 8.5 cm in diameter, solitary, cylindrical, erect, light green to yellowish green with blackish trichomes at emergence, yellowish cream with reddish brown trichomes at maturity; peduncle 10 - 15 cm long, 2.5 - 3.2 cm in diameter, tomentose, reddish brown to brown; microsporophylls 2.33 - 3.0 x 1.14 - 1.90 cm, obconic with a non-recurved distal face and deeply lobate fertile portion, infertile portion 0.45 - 0.65 cm long and rounded with curved horns 0.26 - 0.40 cm long, 0.55 - 0.75 cm and a right angle between the horns. Ovulate strobili 26 - 40 cm long, 11 - 14.5 cm in diameter, solitary, cylindrical, erect, dark green at emergence, dark green with scarce blackish trichomes at maturity, acuminate apex; peduncle 5.0 - 11.0 cm long, 2.5 - 4.0 cm in diameter, erect, tomentose, reddish brown; megasporophylls 153 - 280, 8 - 14 orthostichies with 17 - 20 sporophylls per orthostichy, 1.7 - 2.5 x 2.3 - 3.5 cm, with a prominent distal face, horns straight and 0.38 - 0.50 cm long, 1.35 - 1.60 cm between horns with a right angle between the horns. Seeds 2.5 - 3.5 cm long, 1.9 - 2.5 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
41F0AD37EA8F5A45B4270E4722B73AC4.taxon	distribution	Distribution and habitat. Ceratozamia robusta is widely distributed in Chiapas State, Mexico and northern Guatemala (Fig. 22 B). It occurs in evergreen tropical forest between 400 to 1,300 m on karstic rocks with humus soil.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
41F0AD37EA8F5A45B4270E4722B73AC4.taxon	etymology	Etymology. The epithet is derived from the robust appearance of the species.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
41F0AD37EA8F5A45B4270E4722B73AC4.taxon	discussion	Discussion. Ceratozamia robusta is similar to C. subroseophylla and distinguished by its dark green ovulate strobilus with sparse blackish trichomes on megasporophylls at maturity, prominent distal face, straight horns with a right angle between them, differentiating it easily from C. subroseophylla. The pollen strobilus of C. robusta is the largest in the genus (up to 70 cm long).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
41F0AD37EA8F5A45B4270E4722B73AC4.taxon	materials_examined	Specimens examined. Guatemala. Alta Verapaz: 200 m, 13 Jul 2002, S. Hernandez 757 (MO); 1,800 - 2,000 m, 27 Feb 1939, C. L. Wilson 262 (F); 300 - 500 m, 17 Apr 1942, J. A. Steyermark 45734 (US); 879 m, 15 Mar 2008, M. Veliz et al. 19998 (BIGU). Huehuetenango: 2,000 m, 10 Sep 1942, J. A. Steyermark 52046 (NY, US); 1,150 m, 26 Jul 1943, J. A. Steyermark 49506 (F; MO; NY); 400 m, 27 Jul 1942, J. A. Steyermark 49682 (NY); 900 - 1,300 m, 3 Sep 1942, J. A. Steyermark 51818 (F; NY; US); 900 - 1,000 m, 6 Mar 2009, M. J. M. Christenhusz et al. 5600 (MO); 1,161 m, 21 Dec 2010, L. Velasquez et al. 1566 (BIGU). Mexico. Chiapas: Mun. Berriozabal, 1,129 m, 6 Mar 2014, F. Nicolalde-Morejon et al. 1970 - 1975 (CIB); 1,129 m, 6 Mar 2014, L. Martinez-Dominguez et al. 41 - 47, 49, 50 (CIB), 48 (CIB, MEXU). Mun. Copainala, 1,100 m, M. Gutierrez & T. Acero 240 (CHIP). Mun. Ocosingo, 900 m, 16 Mar 1981, A. P. Vovides & J. I. Calzada 481 (XAL); 14 Apr 1967, D. E. Breedlove 15687 (ENCB); 700 m, 18 Apr 1985, E. Martinez S. 12067 (MEXU); 860 m, 24 Feb 1984, J. Garcia F. 720 (CH, CIB, XAL). Mun. Ocozocoautla de Espinosa, 818 m, 11 Nov 1997, R. Alvarez G. 6 (CHIP). Mun. Osumacinta, 692, 18 Jul 2008, R. Gallegos Ramos 211 (CHIP). Mun. San Fernando, 680 m, 29 Apr 1995, A. P. Vovides & M. A. Perez-Farrera 1266 ª (XAL); 22 Sep 1993, Brigada T. Walters s / n (XAL); 9 Jul 1983, D. W. Stevenson et al. 549 B - F (NY); 1,000 m, 9 Jul 1983, D. W. Stevenson et al. 550 A-C (NY), 24 Apr 1999, M. A. Isidro V. 388 (CHIP); 29 Apr 1995, M. A. Perez-Farrera 293 (MEXU); 12 Jun 2009, N. Martinez-Melendez 2692 (MEXU); 1,200 m, 5 Dec 2000, R. A. Palestina & I. Acosta 2707 (XAL). Mun. Tuxtla Gutierrez, 1,235 m, 19 Mar 1994, M. A. Perez-Farrera 30 (CH, MEXU, XAL); 1,200 m, 1 Nov 1995, M. A. Perez-Farrera 820 (CHIP).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9879DB964EA45F99BF7171F4AF270807.taxon	description	Figs 1 A, 27 D	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9879DB964EA45F99BF7171F4AF270807.taxon	materials_examined	Type. Mexico. Queretaro: Mun. San Joaquin, La Mojonera, 2 km on road San Joaquin-El Aguacate, 1,850 m, 15 Apr 1991, A. P. Vovides & P. Fawcett 1205 ♀ (holotype: XAL! [XAL 0005310]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9879DB964EA45F99BF7171F4AF270807.taxon	description	Description. Stem 8 - 30 cm long, 20 - 35 cm in diameter, epigeous, erect and decumbent. Cataphylls 3 - 4.5 x 2 - 3.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 3 - 40, 60 - 129 cm long, descending, dark brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole 20 - 60 cm long, terete, linear, greenish brown in mature leaves; with 5 - 40 thin prickles, 0.02 - 0.19 cm long. Rachis 40 - 92 cm long, terete, linear, brown in mature leaves, with prickles. Leaflets 26 - 54 pairs, opposite to subopposite, insertion in one plane, linear, longitudinally curved abaxially to planar, basally falcate, papyraceous, flat, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 13 - 32 x 0.6 - 1.5 cm, 0.5 - 1.5 cm between leaflets; articulations 0.3 - 0.7 cm wide, brown. Pollen strobili 11 - 18 cm long, 3.5 - 4.8 cm in diameter, solitary, cylindrical, erect, greenish yellow at emergence, greenish yellow with blackish trichomes at maturity; peduncle 7 - 13 cm long, 1.1 - 1.9 cm in diameter, tomentose, reddish brown to brown; microsporophylls 1.0 - 1.9 x 0.85 - 1.90 cm, discoid with a recurved downward distal face and lobate fertile portion, infertile portion 0.33 - 0.43 cm long and rounded with straight horns 0.06 - 0.20 cm long, 0.30 - 0.71 cm and a right angle between the horns. Ovulate strobili 14 - 19.5 cm long, 5.5 - 8.5 cm in diameter, solitary, cylindrical, erect, yellowish green with brown trichomes at emergence, blue green with blackish trichomes at maturity, apiculate apex; peduncle 3.5 - 7 cm long, 1.2 - 2.5 cm in diameter, erect, tomentose, brown; megasporophylls 72 - 110, 8 - 10 orthostichies with 9 - 11 sporophylls per orthostichy, 3.5 - 5 x 3.8 - 4.5 cm, with a truncate distal face, horns straight and 0.28 - 0.40 cm long, 1.2 - 1.6 cm between horns with a right angle between the horns. Seeds 1.2 - 2.0 cm long, 1.2 - 1.5 cm in diameter, ovate, sarcotesta whitish red when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9879DB964EA45F99BF7171F4AF270807.taxon	distribution	Distribution and habitat. Ceratozamia sabatoi is endemic to the Sierra Gorda mountain range in Mexico, particularly in Queretaro and Hidalgo States (Fig. 22 C), where it occurs in the understory herbaceous layer of the transition zone between oak forest and cloud forest at 1,600 - 1,900 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9879DB964EA45F99BF7171F4AF270807.taxon	etymology	Etymology. The specific epithet honors Sergio Sabato, a distinguished professor at the University of Naples Federico II for his outstanding contributions to knowledge of the biology of Zamiaceae (Vovides et al. 1993).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9879DB964EA45F99BF7171F4AF270807.taxon	discussion	Discussion. Ceratozamia sabatoi differs from all Ceratozamia species by its microsporophylls with a recurved downward distal face. In addition, this species differs from C. kuesteriana by its flat leaflets and descending leaves.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
9879DB964EA45F99BF7171F4AF270807.taxon	materials_examined	Specimens examined. Mexico. Hidalgo: Mun. Jacala de Ledezma, 1,725 m, 16 Aug 2007, A. Frias-Castro et al. 578 (IBUG); 1,500 m, 29 Oct 1946, H. E. Moore, Jr. 1788 (CHAPA). Mun. Zimapan, 2,000 m, 2 Jun 1989, M. Vazquez-Torres & J. P. Sclavo 4035 (CIB); 1,235 m, 22 May 2003, R. Contreras-Medina 55, 56 (XAL); 1,900 m, 13 Sep 1981, R. Fernandez-Nava 6561 (MEXU, MO, XAL). Queretaro: Mun. Cadereyta de Montes, 1,850 m, 15 Apr 1991, A. P. Vovides & K. Norstog 1193 (XAL); A. P. Vovides et al. 1196 - 1199 (XAL), 1203 (MEXU, XAL); 1,850 m, 15 Apr 1991,); A. P. Vovides 1201, 1205 (XAL); 1,924 m, 28 Mar 2015, F. Nicolalde-Morejon et al. 2169, 2170 (CIB); 1,924 m, 28 Mar 2015, L. Martinez-Dominguez et al. 313 - 343 (CIB); 29 Mar 1995, R. Fernandez-Nava s / n (MEXU); 7 Dec 1990, R. Zirahuen-Ortega V. 328 (MEXU). Mun. Landa de Matamoros, 1,439 m, 12 Jan 2001, T. W. Walters 2001 - 05 - A, B (XAL). Mun. Pinal de Amoles, 1,760 m, 17 Sep 2001, E. Carranza G. & I. Silva 6254 (MO); 1,678 m, 29 Mar 2015, F. Nicolalde-Morejon et al. 2171, 2172 (CIB); 1,678 m, 29 Mar 2015, L. Martinez-Dominguez et al. 344 - 372 (CIB); 1,700 m, 11 Dec 1988, Rzedowski s / n (XAL); 1,650 m, 4 Apr 1987, R. Fernandez N. 3819 (ENCB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
D6B2659E41365AE9845F17DCED907646.taxon	description	Figs 9 A, 12 A, 23 D-F	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
D6B2659E41365AE9845F17DCED907646.taxon	materials_examined	Type. Mexico. Chiapas: Mun. Tenejapa, 1,500 m, 16 May 2017, M. A. Pe ́ rez Farrera 3558 ♀ (holotype: HEM [n. v.]; isotypes: MEXU [n. v.], XAL [n. v.]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
D6B2659E41365AE9845F17DCED907646.taxon	description	Description. Stem 8 - 30 cm long, 10 - 20 cm in diameter, semi-hypogeous, erect. Cataphylls 2.5 - 5.8 x 1.3 - 5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 4 - 12 (20), 100 - 232.5 cm long, descending, green or brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole 40 - 134.5 cm long, terete, linear, greenish brown or green in mature leaves; unarmed or with prickles 2 - 28 thin, 0.05 - 0.34 cm long. Rachis 55 - 120 cm long, terete, linear, brown or green in mature leaves, unarmed or with scarce prickles. Leaflets 20 - 35 pairs, opposite to subopposite, insertion in one plane, linear, in general longitudinally planar, not basally falcate (sometimes basally falcate), papyraceous, flat, green with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 22 - 36 x 1.7 - 3.0 cm, 1.4 - 3.5 cm between leaflets; articulations 0.55 - 1.1 cm wide, green. Pollen strobili 15 - 20 cm long, 1.8 - 3.2 cm in diameter, solitary, cylindrical, erect, light green at emergence, green with blackish trichomes at maturity; peduncle 6 - 10 cm long, 0.7 - 1.5 cm in diameter, tomentose, reddish brown to brown; microsporophylls 0.88 - 1.2 x 0.65 - 0.80 cm, discoid with a non-recurved distal face and a lobate fertile portion, infertile portion 0.32 - 0.40 cm long and rounded with straight horns 0.10 - 0.20 cm long, 0.63 - 0.70 cm and an acute angle between the horns. Ovulate strobili 12 - 25 cm long, 7.0 - 8.5 cm in diameter, solitary, cylindrical, erect, yellowish green at emergence, green with scarce blackish trichomes at maturity, acuminate apex; peduncle 5.0 - 11 cm long, 0.9 - 1.1 cm in diameter, erect, tomentose, brown; megasporophylls 35 - 100, 7 - 10 orthostichies with 5 - 10 sporophylls per orthostichy, 0.9 - 1.2 x 3.5 - 4 cm, with a prominent distal face, horns curved to straight and 0.50 - 0.73 cm long, 1.2 - 1.4 cm between horns with a right angle between the horns. Seeds 1.9 - 2.3 cm long, 1.4 - 1.5 cm in diameter, globose, sarcotesta whitish yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
D6B2659E41365AE9845F17DCED907646.taxon	distribution	Distribution and habitat. Ceratozamia sancheziae is endemic to Chiapas State, Mexico (Fig. 22 D), where it occurs in oak forest and pine-oak forest between 1,000 - 1,536 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
D6B2659E41365AE9845F17DCED907646.taxon	etymology	Etymology. This species was named in honor of Maria Ydelia Sanchez-Tinoco, for contributions to our knowledge of the anatomy of Mexican cycad seeds (Gutierrez-Ortega et al. 2021).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
D6B2659E41365AE9845F17DCED907646.taxon	discussion	Discussion. Ceratozamia sancheziae is geographically close to C. robusta, but differs from it by its linear leaflets, leaves at emergence that are light green or reddish brown with a glaucous appearance, and ovulate strobili with curved to straight horns up to 0.73 cm long. Here, we have recircumscribed and clarified the taxonomic identity of Ceratozamia sancheziae. In the description of this species, the authors mentioned Petalcingo as a municipality, but this is a locality that corresponds to Tila municipality in Chiapas (c. f. Gutierrez-Ortega et al. 2021). The distribution range for this species was considered to extend from Tenajapa municipality in Chiapas to the mountain area close to the border with the State of Tabasco (Gutierrez-Ortega et al. 2021). However, we found specimens from Yajalon of Chiapas (M. A. Perez Farrera 1635 from XAL) that do not correspond to the species description (Suppl. material 2). We carried out fieldwork and collected botanical material at the population level in the surroundings of Yajalon, Tila, Altamirano and Tenejapa. Based on the revision of herbarium specimens and the botanical material collected by us, we recircumscribed the populations for Ceratozamia in this region into two species: C. sancheziae and C. zoquorum. One diagnostic character for C. sancheziae is oblanceolate leaflets when juvenile, but we found in the field juveniles with linear leaflets only (Gutierrez-Ortega et al. 2021: 2009; Fig. 23 D). Furthermore, we did not find populations near Yajalon or Tila that correspond to C. sancheziae. The paratypes cited from Yajalon (Mendez Ton 5498, 5722 both MEXU) have some vegetative characters that do not correspond with the species description; however, these could be part of the overall species variation of C. sancheziae. Considering that sympatry is possible in this genus and that there is broad variation in C. zoquorum, research at population level with reproductive structures and molecular variation could clarify what species is represented by these plants collected near Yajalon. In addition, we have extended the range for C. sancheziae to the south of Chiapas (i. e., populations from Altamirano municipality).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
D6B2659E41365AE9845F17DCED907646.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Mun. Altamirano, 1,215 m, 16 Jul 2021, F. Nicolalde-Morejon et al. 3691 - 3697 (CIB); 1,215 m, 16 Jul 2021, L. Martinez-Dominguez 2318 - 2325 (CIB); 1,210 m, 28 Nov 1996, M. A. Perez-Farrera 1468 (CHIP); 1,200 m, 18 May 1993, M. Gonzalez-Espinosa et al., 1973 (CH). Mun. Tenejapa, 1,000 m, 5 Jun 1972, D. E. Breedlove 25506 (MEXU); 1,536 m, 15 Jul 2021, F. Nicolalde-Morejon et al. 3683 - 3690 (CIB); 1,536 m, 15 Jul 2021, L. Martinez-Dominguez et al. 2310 - 2317 (CIB). Mun. Yajalon, 1,434 m, 26 Aug 2015, F. Hernandez-Najarro 4382 (CHIP).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
40FB50ED8EC05A09B4FC8A8E6D0AC781.taxon	description	Figs 1 B, 27 E	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
40FB50ED8EC05A09B4FC8A8E6D0AC781.taxon	materials_examined	Type. Mexico. Chiapas: Mun. Berrioza ́ bal, road Berrioza ́ bal-El Cairo, 15 Oct 2004, M. A. Pe ́ rez Farrera 3030 ♀ (holotype: HEM! [HEM 020981]; isotypes: XAL! [XAL 0005415], MEXU [n. v.]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
40FB50ED8EC05A09B4FC8A8E6D0AC781.taxon	description	Description. Stem 10 - 50 cm long, 11 - 13 cm in diameter, semi-hypogeous, erect and decumbent. Cataphylls 2 - 4.5 x 1.7 - 4 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, with apex partiality glabrous at maturity, apex acuminate. Leaves 2 - 3, 50.5 - 79.3 cm long, descending, light green and glaucous at emergence, with whitish gray trichomes, green and glabrous at maturity. Petiole 23.5 - 45 cm long, terete, linear, yellowish green green in mature leaves; with 4 - 15 thin prickles, 0.05 - 0.15 cm long or unarmed. Rachis 23 - 40 cm long, terete, linear, yellowish green green at emergence, yellowish green in mature leaves, unarmed to armed with prickles. Leaflets 6 - 12 pairs, opposite to subopposite, insertion in one plane, oblong, longitudinally curved abaxially to planar, not basally falcate to basally falcate, coriaceous, flat, green with adaxial and abaxial sides glaucous, distal end with entire margins, acuminate and asymmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 17.4 - 30.6 x 4.2 - 6.5 cm, 3.5 - 7.1 cm between leaflets; articulations 0.7 - 1.1 cm wide, yellow. Pollen strobili 15 - 20 cm long, 1.5 - 3.0 cm in diameter, solitary, cylindrical, erect, green with blackish trichomes at emergence, yellow-cream with blackish at maturity; peduncle 2 - 4 cm long, 1.0 - 1.3 cm in diameter, scarce pubescent, reddish brown to brown; microsporophylls 1.2 - 2 x 0.5 - 1 cm, discoid with a non-recurved distal face and a lobate fertile portion, infertile portion 0.34 - 0.37 cm long and linear with straight horns 0.20 - 0.25 cm long, 0.50 - 0.65 cm and an acute angle between the horns. Ovulate strobili 12 - 20 cm long, 7 - 9 cm in diameter, solitary, cylindrical, erect, green with blackish trichomes at emergence, green with scarce blackish trichomes at maturity, acute apex; peduncle 2 - 3 cm long, 0.9 - 1.3 cm in diameter, erect, tomentose, light brown; megasporophylls 21 - 56, 7 - 8 orthostichies with 3 - 7 sporophylls per orthostichy, 1.3 - 2.2 x 1.5 - 2.2 cm, with a prominent distal face, horns straight and 0.60 - 0.80 cm long, 1.65 - 1.90 cm between horns with an acute angle between the horns. Seeds 2.2 - 2.4 cm long, 1.8 - 2.0 cm in diameter, ovate, sarcotesta whitish pink when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
40FB50ED8EC05A09B4FC8A8E6D0AC781.taxon	distribution	Distribution and habitat. Ceratozamia santillanii is endemic to the northern highlands of Chiapas State, Mexico and only known from the type locality in the municipality of Berrioza ́ bal (Fig. 24 A), where it was collected in evergreen tropical rain forest on karstic rocks at 800 - 900 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
40FB50ED8EC05A09B4FC8A8E6D0AC781.taxon	etymology	Etymology. The specific epithet was assigned in honor of Professor Trinidad Alema ́ n Santilla ́ n in recognition of his academic accomplishments in the training of young biologists in botany and ecology (Perez-Farrera et al. 2009).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
40FB50ED8EC05A09B4FC8A8E6D0AC781.taxon	discussion	Discussion. Ceratozamia santillanii belongs to a cryptic taxonomic group with C. zoquorum and C. becerrae, which is characterized by oblong and coriaceous leaflets with an acuminate and asymmetric apex. C. santillanii is easily distinguished from the other taxa in this complex with internal transcribed spacer region of nuclear ribosomal (nrITS) (Martinez-Dominguez et al. 2017 c), and peduncle of ovulate strobilus 3 cm long or shorter.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
40FB50ED8EC05A09B4FC8A8E6D0AC781.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Known only from the type locality. 15 Oct 2004, M. A. Perez-Farrera 2944 (HEM).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C00E8546A3555D3CA569ECB311880B43.taxon	description	Figs 12 D, 25, 26	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C00E8546A3555D3CA569ECB311880B43.taxon	materials_examined	Type. Mexico. Veracruz: Mun. Santiago Tuxtla, hill in front Sinapan, 425 m, 15 Jul 2014, L. Marti ́ nez-Domi ́ nguez 158 ♀ (holotype: CIB! [acc. # 16893 UV]; isotypes: MEXU! [MEXU 01446538 - MEXU 01446542], NY!).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C00E8546A3555D3CA569ECB311880B43.taxon	description	Description. Stem 30 - 250 (500) cm long, 18 - 45 cm in diameter, epigeous, erect and decumbent. Cataphylls 5 - 7 x 2 - 2.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 10 - 83, 72 - 370 cm long, ascending, yellowish brown at emergence, with whitish gray trichomes, glabrous at maturity. Petiole 50 - 150 cm long, terete, linear, greenish brown or dark brown at emergence, green in mature leaves; with 25 - 55 robust prickles, 0.40 - 0.85 cm long. Rachis 80 - 255 cm long, terete, linear, greenish brown or dark brown at emergence, green in mature leaves, with prickles. Leaflets 23 - 48 pairs, opposite to subopposite, insertion in one plane, lanceolate, abaxially curved longitudinally along distal half, not basally falcate, papyraceous, flat, dark green with adaxial side glabrous and abaxial side glaucous, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 18 - 45 x 2.5 - 4 cm, 1.7 - 6.5 cm between leaflets; articulations 0.7 - 1.9 cm wide, brown in young leaves and green in mature leaves. Pollen strobili 15 - 30 cm long, 3.1 - 5.4 cm in diameter, solitary, cylindrical, erect, green with reddish trichomes at emergence, greenish yellow at maturity with dark brown trichomes; peduncle 9.5 - 19 cm long, 1.5 - 2.7 cm in diameter, tomentose, reddish brown; microsporophylls 1.47 - 2.8 x 1.01 - 1.4 cm, obconic with a non-recurved distal face and lobate fertile portion, infertile portion 0.49 - 0.65 cm long and rounded with straight horns 0.20 - 0.30 cm long, 0.48 - 0.53 cm and an acute angle between the horns. Ovulate strobili 15.5 - 40 cm long, 7 - 11.6 cm in diameter, solitary, cylindrical, erect, yellow with reddish to purple trichomes at emergence, green and megasporophylls with base pale pink and dark brown to reddish brown trichomes at maturity, mucronate apex; peduncle 9.8 - 17.5 cm long, 1.8 - 2.3 cm in diameter, erect, tomentose, light brown; megasporophylls 110 - 210, 9 - 12 orthostichies with 11 - 20 sporophylls orthostichy, 1.8 - 2.5 x 2.5 - 4.47 cm, with a prominent distal face, horns straight 0.34 - 1.0 cm long and 0.70 - 1.40 cm between horns with a right angle between the horns. Seeds 2.9 - 3.8 cm long, 0.80 - 1.75 cm in diameter ovate, sarcotesta whitish pink when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C00E8546A3555D3CA569ECB311880B43.taxon	distribution	Distribution and habitat. Ceratozamia subroseophylla occurs from southeastern Veracruz State to Tabasco State, Mexico including the montane zone of the region Santiago Tuxtla and Uxpanapa (Fig. 24 B). It occurs on soils of volcanic origin and karstic rocks in evergreen tropical rain forest between 111 and 1,050 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C00E8546A3555D3CA569ECB311880B43.taxon	etymology	Etymology. The specific epithet efers to the rosaceous (i. e., pale pink) color at the base of megasporophylls.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C00E8546A3555D3CA569ECB311880B43.taxon	discussion	Discussion. Ceratozamia subroseophylla is distinguished from C. leptoceras and C. oliversacksii by its green-brownish petiole and rachis with abundant and robust prickles, and its lanceolate and papyraceous leaflets (Fig. 25). In addition, C. subroseophylla has affinity with C. robusta, but it is easily identified by the ovulate strobilus which has mucronate apex, rosaceous base of megasporophylls rosaceous with dark brown to reddish brown trichomes at maturity (Fig. 26). Populations from Uxpanapa (Veracruz) were recently described as Ceratozamia dominguezii, however, these fall within the range of variation of C. subroseophylla as circumscribed here. Our circumscription is based on comparative morphology, both vegetative and reproductive structures, and phenology (Figs 25, 26).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C00E8546A3555D3CA569ECB311880B43.taxon	materials_examined	Specimens examined. Mexico. Tabasco: Mun. Macuspana, 185 m, 13 Jul 2021, F. Nicolalde-Morejon et al. 3674 - 3680 (CIB); 185 m, 13 Jul 2021, L. Martinez-Dominguez et al. 2295 - 2300 (CIB), 190 m, 13 Jul 2021, L. Martinez-Dominguez et al. 2301 (CIB, MEXU). Veracruz: Mun. Catemaco, 1953, H. Bravo 26 (MEXU); 18 May 1995, M. A. Garci ́ a B. et al. 813 (XAL). Mun. Hidalgotitlan, 150 m, 16 Jan 1975, Brigada Vazquez 1760 (MO, XAL); 140 m, 16 Dec 1974, J. Rees 1655 (XAL); 16 Apr 1975, M. Vazquez-Torres 1760 (MEXU). Mun. Hueyapan de Ocampo, 400 m, 14 May 2000, Leonti 542 (MEXU). Mun. Jesus Carranza, 100 m, 19 Feb 2009, D. Jimeno-Sevilla 1045 (MEXU, MO, XAL); 131 m, 27 Sep 2020, L. Martinez-Dominguez et al. 2040 (CIB); M. Vazquez-Torres 2430 (CHAPA, ENCB, MEXU, NY, SLPM, XAL). Mun. Mecayapan, 950 m, 26 Jan 1992, M. Va ́ zquez-Torres et al. 4122 (CIB). Mun. Santiago Tuxtla, 500 m, 5 Jul 1983, D. W. Stevenson et al. 539 A-L (NY), 540 F-H (NY); 420 m, 28 Mar 2014, F. Nicolalde-Morejo ́ n & L. Marti ́ nez-Domi ́ nguez 1985 - 1987 (CIB); 250 - 500 m, 30 Sep 1983, J. F. Ortega O. et al. 358 (XAL); 420 m, 28 Mar 2014, L. Marti ́ nez-Domi ́ nguez & F. Nicolalde-Morejo ́ n 78, 79 (NY), 80, 81 (XAL), 82 - 87 (CIB); 420 m, 7 Jun 2014, L. Marti ́ nez-Domi ́ nguez & F. Nicolalde-Morejo ́ n 129 (CIB); 425 m, 15 Jul 2014, L. Marti ́ nez-Domi ́ nguez 136 - 157, 159 (CIB); 500 m, 12 May 1965, M. Sousa 2420 (MEXU); 23 Aug 1962, R. F. Andrle 64 (US); 497 m, 17 Jan 2001, T. W. Walters TW- 2001 - 17 (MEXU, XAL). Mun. Soteapan, 21 Jun 1963, G. N. Ross 57 (US); 1,050 m, 17 Mar 1968, M. Sousa 3645 (MEXU); 500 m, 18 May 1986, M. Va ́ zquez-Torres et al. 3579 (CIB). Mun. Tatahuicapan de Jua ́ rez, 849 m, C. I. Carvajal-Herna ́ ndez & M. Jua ́ rez F. 796 (CIB). Mun. Uxpanapa, 131 m, 27 Sep 2020, L. Marti ́ nez-Domi ́ nguez et al. 2040 (CIB, MEXU); 111 m, 27 Sep 2020, L. Marti ́ nez-Domi ́ nguez et al. 2041 (CIB), 2042 (CIB, MEXU), 2043 - 2048 (CIB); 111 m, 11 Jun 2021, L. Marti ́ nez-Domi ́ nguez et al. 2273 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C0EA45994A565E5299031B2A4F57ABDF.taxon	description	Fig. 27 F	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C0EA45994A565E5299031B2A4F57ABDF.taxon	materials_examined	Type. Based on Ceratozamia mexicana Brongn. var. tenuis Dyer, Biol. Cent. - Amer., Bot. 3: 193. 1884.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C0EA45994A565E5299031B2A4F57ABDF.taxon	description	Description. Stem 20 - 100 cm long, 30 - 45 cm in diameter, epigeous, erect and decumbent. Cataphylls 2 - 6 x 2 - 5.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 6 - 56, 85 - 225 cm long, ascending, dark green at emergence with brown trichomes, glabrous at maturity. Petiole 30 - 93 cm long, terete, linear, green in mature leaves; with 10 - 55 thin prickles, 0.20 - 0.49 cm long. Rachis 56 - 154 cm long, terete, linear, green in mature leaves, with prickles. Leaflets 30 - 56 pairs, opposite to subopposite, insertion in one plane, linear, longitudinally curved abaxially to planar, basally falcate, papyraceous, involute, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at apex, attenuate at base, with conspicuous and light-green veins; median leaflets 23 - 50.5 x 1 - 2.1 cm, 0.3 - 2.5 cm between leaflets; articulations 0.4 - 1.4 cm wide, green. Pollen strobili 25 - 50 cm long, 4 - 8 cm in diameter, solitary, cylindrical, erect, greenish yellow at emergence, greenish yellow with blackish trichomes at maturity; peduncle 3.7 - 22 cm long, 1.2 - 2.5 cm in diameter, tomentose, reddish brown to light-brown; microsporophylls 1.7 - 2.7 x 1.2 - 1.9 cm, obconic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.50 - 0.65 cm long and orbicular with recurved horns 0.25 - 0.46 cm long, 0.35 - 0.65 cm between the horns and a right angle between the horns. Ovulate strobili 22 - 35 cm long, (7.6) 10 - 14 cm in diameter, solitary, cylindrical, erect, dark green with blackish trichomes at emergence, dark green with blackish trichomes at maturity, acuminate apex; peduncle 8 - 23 cm long, 1.5 - 2.4 cm in diameter, erect or pendulous, tomentose, brown to reddish brown; megasporophylls (48) 80 - 196, 7 - 16 orthostichies with 6 - 14 sporophylls per orthostichy, 2.1 - 3.1 x 3.0 - 5.0 cm, with a prominent distal face, horns curved and 0.32 - 0.80 cm long, 0.80 - 1.60 cm between horns with a right angle between the horns. Seeds 2.5 - 3 cm long, 1.3 - 1.8 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C0EA45994A565E5299031B2A4F57ABDF.taxon	distribution	Distribution and habitat. Ceratozamia tenuis is endemic to the central montane region in Veracruz State, Mexico (Fig. 24 C), where it occurs in cloud forest at 1,200 - 1,920 m elevation on volcanic soils with basaltic rocks.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C0EA45994A565E5299031B2A4F57ABDF.taxon	etymology	Etymology. The specific epithet is derived from its thin leaflets.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C0EA45994A565E5299031B2A4F57ABDF.taxon	discussion	Discussion. Ceratozamia tenuis is characterized by a petiole with thin prickles, and linear leaflets that are papyraceous and involute with a symmetric apex. The ovulate strobilus is dark green with blackish trichomes at maturity, a prominent distal face, and a right angle between the horns.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
C0EA45994A565E5299031B2A4F57ABDF.taxon	materials_examined	Specimens examined. Mexico. Veracruz: Mun. Banderilla, 1,450 m, 21 Apr 2017, L. Martinez-Dominguez et al. 1000 (CIB). Mun. Chiconquiaco, 1,800 m, 26 Nov 1974, D. Jimeno-Sevilla 754 (XAL); 1,916 m, 27 Sep 2016, F. Nicolalde-Morejon et al. 2456 - 2464 (CIB); 1,800 m, 26 Nov 1974, J. Rees 1625 (XAL), 1626 (MEXU, XAL); 1,916 m, 27 Sep 2016, L. Martinez-Dominguez et al. 971, 973 - 981 (CIB), 972 (CIB, MEXU); 13 Apr 1967, R. Fernandez-Nava 385 A (MEXU). Mun. Coacoatzintla, 8 Mar 1985, F. Vazquez B. 2275 (XAL); 1,400 m, 7 Jan 1977, G. Castillo-Campos 118 (XAL); 1,550 m, 3 Jun 2005, L. H. Bojorquez-Galvan & A. M. Zapata-Aquino 1484, 1485 (CIB); 1,435 m, 9 Jan 2015, L. Martinez-Dominguez et al. 165 - 184 (CIB); 1, 435 m, 18 Mar 2015, L. Martinez-Dominguez et al. 273 - 282 (CIB); 1,540 m, 9 Feb 2016, L. Martinez-Dominguez et al. 759 (CIB). Mun. Jilotepec, 1,300 m, 29 Feb 1980, A. P. Vovides 470 (XAL), 471 (IBUG, MEXU, XAL); 1,300 m, 14 Apr 1982, 735 (XAL); 1,385 m, 15 Nov 1978, E. Estrada et al. 757 (MEXU); 1,316 m, 22 Aug 2014, F. Nicolalde-Morejon & L. Martinez-Dominguez 2067 - 2086 (CIB); 1,350 m, 7 Dec 1970, F. Ventura A. 2936 (ENCB); 1,300 m, 22 Jan 1971, F. Ventura A. 3014 (ENCB); 1,300 m, 18 Nov 1974, J. Rees 1620 (XAL); 1,363 m, 29 May 2015, L. Martinez-Dominguez et al. 573 - 583 (CIB); 1,250 m, 23 Aug 1973, M. G. Zola 657 (XAL), 667 (MEXU, XAL); 1,250 m, 23 Aug 1975, R. Ortega J. 525 (XAL); Mar 2001, S. Avendano 5395 (MEXU). Mun. Tepetlan, 1,420 m, 12 Jan 2013, F. Nicolalde-Morejon et al. 1691 - 1710 (CIB); 1,418 m, 21 Jun 2014, F. Nicolalde-Morejon & L. Martinez-Dominguez 2001 - 2004 (CIB); 1,418 m, 22 Aug 2014, F. Nicolalde-Morejon & L. Martinez-Dominguez 2047 - 2066 (CIB); 1,662 m, 10 Apr 2015, F. Nicolalde-Morejon & L. Martinez-Dominguez 2217 - 2226 (CIB); 1,421 m, 26 Jul 2014, L. Martinez-Dominguez et al. 160 (CIB); 22 Mar 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 283 - 293 (CIB); 1,662 m, 10 Apr 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 545 - 555 (CIB); 2 Jul 2010, M. Vazquez-Torres et al. 9215 (CIB). Mun. Tlacolulan, 1,540 m, 16 Jun 2017, F. Nicolalde-Morejon et al. 2516 (CIB); 1,540 m, 16 Jun 2017, L. Martinez-Dominguez et al. 1041, 1042 (CIB). Mun. Xalapa, 1,270 m, 18 Oct 2016, F. Nicolalde-Morejon et al. 2466 - 2468 (CIB); 1,270 m, 18 Oct 2016, L. Martinez-Dominguez et al. 985, 987 (CIB), 986 (CIB, MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
7DCCF1E75A3953AFA5C2A3ECF6D7FABA.taxon	description	Figs 27 G, 28 B	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
7DCCF1E75A3953AFA5C2A3ECF6D7FABA.taxon	materials_examined	Type. Mexico. Puebla: Mun. Jonotla, 600 m, 9 Jun 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 618 ♀ (holotype: CIB! [acc. # 16735 UV]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
7DCCF1E75A3953AFA5C2A3ECF6D7FABA.taxon	description	Description. Stem 10 - 45 cm long, 10 - 25 cm in diameter, epigeous, erect and decumbent. Cataphylls 2 - 5 x 1.2 - 2.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at maturity, apex acuminate. Leaves 10 - 63, 100 - 265 cm long, descending, brown at emergence, with brown trichomes, glabrous at maturity. Petiole 30 - 80 cm long, terete, straight, green in mature leaves; with 10 - 40 thin prickles, 0.05 - 0.25 cm long. Rachis 85 - 185 cm long, terete, straight, green in adult leaves, with prickles. Leaflets 11 - 33 pairs, opposite to subopposite, insertion in one plane, oblong, longitudinally planar, not basally falcate to occasionally falcate, papyraceous, flat, green with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 17 - 40 x 2.7 - 4.2 cm, 2 - 5.6 cm between leaflets; articulations 0.5 - 1.3 cm wide, green. Pollen strobili 28 - 31 cm long, 5.0 - 6.0 cm in diameter, generally solitary (up to 2), cylindrical, erect, greenish yellow at emergence, yellow with brown trichomes at maturity; peduncle 9 - 12 cm long, 1.5 - 2 cm in diameter, tomentose, light brown; microsporophylls 1.5 - 2.7 x 0.9 - 2.0 cm, obconic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.39 - 0.57 cm long and rounded with straight horns 0.19 - 0.25 cm long, 0.52 - 0.69 cm and a right angle between the horns. Ovulate strobili 20.5 - 28.7 cm long, 8.4 - 9.3 cm in diameter, solitary, cylindrical, erect, light green and glaucous, with orange to light brown trichomes at emergence, green with yellowish brown trichomes at maturity, acuminate apex; peduncle 10 - 11.2 cm long, 1.5 - 2.4 cm in diameter, erect or pendulous, tomentose, light brown; megasporophylls 64 - 120, 8 - 10 orthostichies with 8 - 13 sporophylls per orthostichy, 1.6 - 2.3 x 2.6 - 3.6 cm, with a prominent distal face, horns straight and 0.55 - 0.80 cm long, 1.45 - 1.80 cm between horns with a right angle between the horns. Seeds 2.5 - 3.5 (4) cm long, 0.88 - 1.6 cm in diameter, ovate, sarcotesta whitish red when immature, cream to light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
7DCCF1E75A3953AFA5C2A3ECF6D7FABA.taxon	distribution	Distribution and habitat. Ceratozamia totonacorum occurs in the Sierra Norte de Puebla and the mountain region in Hidalgo and Veracruz States, Mexico (Fig. 24 D). It occurs in cloud forest and the transition zone between evergreen tropical forest and cloud forest on rocky outcrops in exposed walls up to 80 m tall at 600 - 1,800 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
7DCCF1E75A3953AFA5C2A3ECF6D7FABA.taxon	etymology	Etymology. The specific epithet is in reference to the Totonaco ethnic group of Santiago Ecatla ́ n in Sierra Norte of Puebla, Mexico.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
7DCCF1E75A3953AFA5C2A3ECF6D7FABA.taxon	discussion	Discussion. Ceratozamia totonacorum is distinguished from C. delucana by its brown leaves at emergence, but this color disappears in the mature leaves which become green; besides, the leaves are descending. The ovulate strobilus is yellowish green with brown trichomes.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
7DCCF1E75A3953AFA5C2A3ECF6D7FABA.taxon	materials_examined	Specimens examined. Mexico. Hidalgo: Mun. Huehuetla, 1,150 m, 2 Jun 1976, A. P. Vovides 23 (XAL). Puebla: Mun. Atlequizayan, 1 Apr 2014, 867 m, L. Caamano Onofre & A. B. Ceron Carpio 4995 (XAL). Mun. Cuetzalan del Progreso, 1,470 m, 1 Nov 2017, L. Martinez-Dominguez et al. 1180 - 1189 (CIB). Mun. Jonotla, 760 m, 13 Feb 2014, F. Nicolalde-Morejon et al. 1948 (CIB); 600 m, 13 Feb 2014, F. Nicolalde-Morejon et al. 1949 - 1955 (CIB), 1956 (MEXU), 1957 (NY), 1958 - 1965 (CIB); 1,006 m, 14 Feb 2014, F. Nicolalde-Morejon et al. 1966, 1967 (CIB); 600 m, 9 Jun 2015, L. Martinez-Dominguez & F. Nicolalde-Morejon 619 (CIB). Mun. Pahuatlan, 1,800 m, 12 Jan 1987, G. Toriz et al. 226 (MEXU). Mun. Tlapacoya, 1,010 m, 14 Feb 1985, E. Meza P. 14 (XAL). Mun. Zacapoaxtla, 1, 365 m, 30 May 2014, L. Camano-Onofre 5329 (XAL). Veracruz: Mun. Tlachichilco, 1,300 m, 12 Sep 2001, A. Rincon G. et al. 2584 (XAL), 2585 (MEXU, XAL).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AE47E89386015B429736BE27E41194B1.taxon	description	Fig. 27 H	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AE47E89386015B429736BE27E41194B1.taxon	materials_examined	Type. Mexico. Chiapas: Mun. La Concordia, Between Finca Santa Cruz and El Puente, Rancho Las Cabanas, 1,156 m, 19 Jul 2001, M. A. Perez-Farrera 2620 a ♀ (holotype: HEM!; isotypes: CHIP [n. v.], MEXU [n. v.], MO [n. v.], XAL [n. v.]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AE47E89386015B429736BE27E41194B1.taxon	description	Description. Stem 50 - 80 cm long, 20 - 30 cm in diameter, epigeous, erect and decumbent. Cataphylls 5 - 7.5 x 2.7 - 5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 3 - 18, 97 - 238 cm long, ascending, reddish brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole terete, linear, 40 - 164 cm long, green in adult leaves, with 15 - 55 thin prickles, 0.14 - 0.38 cm long. Rachis terete, linear, 54 - 153 cm long, green in mature leaves, with prickles. Leaflets 30 - 85, opposite to subopposite, insertion in one plane, lanceolate, mostly longitudinally planar, generally basally falcate, papyraceous, flat, green with adaxial side glabrous and abaxial side glaucous, distal end with entire margins, acuminate and symmetrical apex, attenuate at base, with conspicuous and light-green veins; median leaflets 20 - 45 x 0.7 - 1.4 cm, 0.3 - 2.4 cm between leaflets; articulations 0.2 - 0.8 cm wide, green. Pollen strobili 15 - 45 cm long, 3.5 - 5 cm in diameter, solitary, cylindrical, erect, yellowish green with reddish brown trichomes at emergence, yellowish cream with reddish brown trichomes at maturity; peduncle 6 - 9.5 cm long, 1.6 - 1.9 cm in diameter, tomentose, reddish brown to brown; microsporophylls 1.2 - 1.5 x 0.8 - 1.04 cm, obconic with a non-recurved distal face and lobate fertile portion, infertile portion 0.37 - 0.50 cm long and rounded with straight horns 0.30 - 0.42 cm long, 0.52 - 0.84 cm with a right angle between the horns. Ovulate strobili 26 - 40 cm long, 7.1 - 9.6 cm in diameter, solitary, cylindrical, erect, greyish green with abundant reddish brown trichomes at emergence, green with abundant blackish trichomes at maturity, acuminate apex; peduncle 7 - 15 cm long, 1.7 - 2.2 cm in diameter, erect or pendulous, pubescent, brown; megasporophylls 60 - 70, 6 - 8 orthostichies with 7 - 12 sporophylls per row, 3.8 - 4.5 x 4 - 5 cm, with a prominent distal face, horns straight and 0.60 - 0.80 cm long, 0.99 - 1.40 cm between horns with an obtuse angle between the horns. Seeds 2.2 - 2.7 cm long, 1.3 - 1.7 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AE47E89386015B429736BE27E41194B1.taxon	distribution	Distribution and habitat. Ceratozamia vovidesii is distributed along the Sierra Madre of Chiapas State in Mexico to Guatemala. It occurs on karstic rocks in cloud forest between 800 and 1,850 m elevation (Fig. 29 A).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AE47E89386015B429736BE27E41194B1.taxon	etymology	Etymology. The specific epithet honors Andrew P. Vovides in recognition of his systematic and ecological studies and efforts in the conservation of Mexican cycads, including the creation of the Mexican National Cycad Collection at the Francisco Javier Clavijero Botanic Garden in Xalapa (Veracruz, Mexico) (Perez-Farrera et al. 2007).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AE47E89386015B429736BE27E41194B1.taxon	discussion	Discussion. Ceratozamia vovidesii shares several vegetative characteristics with C. mirandae. However, there are differences in their pollen strobili; C. vovidesii has obconic microsporophylls with rounded infertile portions, whereas those of C. mirandae are elliptic with a linear infertile portion. There is some confusion concerning the holotype for C. vovidesii. In 2012 the gathering Perez-Farrera 2620 a also was used as the holotype in the protologue of Zamia grijalvensis Perez-Farr., Vovides & Mart. - Camilo (Perez-Farrera et al. 2012). The specimen, Perez-Farrera 2620 a is clearly a Ceratozamia and not a Zamia and corresponds to the protologue of C. vovidesii. Another specimen, Perez-Farrera 3026, is labelled as the holotype for Z. grijalvensis and it clearly matches the protologue of that taxon. We are treating this miscitation of Perez-Farrera 2620 a in the protologue of Z. grijalvensis as a typographical error, which is now corrected.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
AE47E89386015B429736BE27E41194B1.taxon	materials_examined	Specimens examined. Guatemala. Huehuetenango: 900 - 1,300 m, 3 Sep 1942, J. A. Steyermark 51818 (NY; US); 1,630 m, 9 Jul 2006, M. Veliz & V. Davila 17042, 17043, 17044 (BIGU); 30 May 1906, O. F. Cook 51 (US); 1,629 m, 10 Jul 2006, V. Davila & M. Veliz 1050 (BIGU); 1,622 m, 10. Jul 2006, V. Davila & M. Veliz 1052, 1053 (BIGU). Mexico. Chiapas: Mun. Angel Albino Corzo, 730 m, 23 Jan 1968, Alush Shilom Ton 3554 (ENCB); 14 Dec 1980, 1,380 m, D. E. Breedlove 48678 (NY); 800 - 1,000 m, 8 Nov 1945, E. H. Xolocotzi & A. J. Sharp 402 (CHAPA, ENCB, MEXU); 1,000 m, 2 Jun 1987, E. M. Martinez S. et al. 21586 (MEXU); 1,819 m, 13 Aug 2009, H. Gomez-Dominguez 2316 (MEXU); 1,650 m, 18 May 1982, J. I. Calzada et al. 9131 (XAL); 5 Mar 1989, U. Bachem C. & R. Rojas 405 (CHIP). Mun. Jaltenango de la Paz, Jun 1995, E. Matuda s / n (MEXU); 1,500 m, 23 Jun 1990, M. Heath & A. Long 1287 (CHIP); 25 Feb 1995, Miranda 7042 (MEXU). Mun. La Concordia, 1,000 m, 5 Jun 1988, E. Palacios E. 1050 (CHIP); 1,840 m, 26 Jun 2018, F. Nicolalde-Morejon et al. 2864 - 2875 (CIB); 1,840 m, 26 Jun 2018, L. Martinez-Dominguez et al. 1439 - 1451 (CIB); 1,156 m, 19 Sep 2001, M. A. Perez-Farrera 2621 (XAL); 1,600 m, M. A. Perez-Farrera s / n (CHIP); 1,120 m, 17 Jun 2014, M. G. Diaz M. 961 (CHIP); 24 Mar 2001, R. Martinez-Camilo 54 (CHIP); 1,100 m, 11 Jun 1988, T. G. Cabrera-Cachon 74 (CHIP); 1,700 m, 1 Jun 1989, U. Bachem C. & R. Rojas 795 (CHIP). Mun. Mapastepec, 1,750 m, 13 May 1982, J. I. Calzada et al. 8874 (IBUG, MEXU, MO, XAL). Mun. Siltepec, 28 Feb 2000, O. Farrera S. 1958 (CHIP).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
209342C85E725D53B4F2DAF76CB05429.taxon	description	Fig. 30	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
209342C85E725D53B4F2DAF76CB05429.taxon	materials_examined	Type. Mexico. Oaxaca: Vicinity of Metates, south of Valle Nacional, 628 m, 10 May 1995, J. Chemnick & T. Gregory 5 (holotype: HNT [n. v.]; isotypes: FTG!, XALU [n. v.]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
209342C85E725D53B4F2DAF76CB05429.taxon	description	Description. Stem 20 - 80 cm long, 18 - 30 cm in diameter, epigeous, decumbent. Cataphylls 2 - 5 x 2 - 5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 3 - 10, 129 - 250 cm long, descending, light green and glaucous at emergence with whitish gray trichomes, glabrous at maturity. Petiole 60 - 140 cm long, terete, linear, green in mature leaves; with 3 - 22 thin prickles, 0.06 - 0.39 cm long. Rachis 70 - 160 cm long, terete, linear, greenish in mature leaves, unarmed. Leaflets 20 - 48 pairs, opposite to subopposite, insertion in one plane, lanceolate, generally longitudinally planar, not basally falcate, papyraceous, flat, green with adaxial and abaxial sides glabrous, distal end with entire margins, acuminate and symmetrical at the apex, attenuate at base, with conspicuous and green-light veins; median leaflets 25 - 38.5 x 2.0 - 3.7 cm, 1.6 - 3.1 cm between leaflets; articulations 0.5 - 1.4 cm wide, green. Pollen strobili 20 - 30 cm long, 3 - 5 cm in diameter, generally solitary (1 - 2), cylindrical, erect, greenish at emergence with reddish trichomes, greenish yellow with reddish brown trichomes at maturity; peduncle 15 - 25 cm long, 1.2 - 1.9 cm in diameter, glabrous or with trichomes scarce reddish brown to brown; microsporophylls 1.5 - 3.0 x 0.8 - 1.5 cm, elliptic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.40 - 0.50 cm long and linear with straight horns 0.38 - 0.50 cm long, 0.50 - 1.0 cm and a right angle between the horns. Ovulate strobili 14 - 20 cm long, 7 - 10 cm in diameter, solitary, cylindrical, erect, yellowish green with abundant blackish trichomes at emergence, green with brown to blackish trichomes at maturity, apiculate apex; peduncle 1 - 4 cm long, 1.0 - 1.8 cm in diameter, erect, glabrous or with trichomes scarce, reddish brown; megasporophylls 24 - 40, 4 - 5 orthostichies with 5 - 8 sporophylls per orthostichy, 1.7 - 2.2 x 4.5 - 5.1 cm, with a truncate distal face, horns straight and 0.98 - 1.4 cm long, 0.95 - 1.3 cm between horns and an acute angle between the horns. Seeds 2.1 - 2.7 cm long, 1.4 - 1.8 cm in diameter, ovate, sarcotesta whitish yellow to yellow when immature, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
209342C85E725D53B4F2DAF76CB05429.taxon	distribution	Distribution and habitat. Ceratozamia whitelockiana is endemic to the Sierra Norte of Oaxaca (Mexico), between 500 to 1,800 m in La Chinantla area (Fig. 29 B). It occurs in the elevational gradient of evergreen tropical forest with Quercus sp. and cloud forest with Oreomunnea mexicana (Standl.) J. - F. Leroy on karstic rocks.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
209342C85E725D53B4F2DAF76CB05429.taxon	etymology	Etymology. This species was named in honor of Loran Whitelock for his contributions to cycad biology (Chemnick and Gregory 1996).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
209342C85E725D53B4F2DAF76CB05429.taxon	discussion	Discussion. Ceratozamia whitelockiana is similar to C. mixeorum in leaf morphology; the only difference is the sparse prickles in C. whitelockiana (between 3 to 22, in number), whereas C. mixeorum has more than 28 prickles. Additionally, the length of the petiole is very long in relation to the total size of the leaf in C. whitelockiana. In reproductive structures, the ovulate strobili in C. whitelockiana have a long peduncle 12 - 23 cm and the fertile portion has 4 - 5 orthostichies with 5 - 8 sporophylls per orthostichy and the pollen strobili have a long peduncle that is the same size as the fertile part or longer (Fig. 30 A, C). In contrast, C. mixeorum has ovulate strobili with a short peduncle from 1 - 4 cm long and pollen strobili with peduncles shorter than the fertile area.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
209342C85E725D53B4F2DAF76CB05429.taxon	materials_examined	Specimens examined. Mexico. Oaxaca: 660 m, 29 Jun 1977, T. B. Croat 39751 (MO). Mun. Ixtlan de Juarez, 1,640 m, 29 Jan 1998, Y. Arellanes C. et al. 283 (SERO); 1,640 m, 21 Aug 1998, Y. Arellanes C. et al. 413 (MEXU, MO, SERO). Mun. San Felipe Usila, 24 Oct 1994, P. Osorio H. 312 (MEXU). Mun. San Juan Bautista Valle Nacional, 650 m, 24 Sep 2020, F. Nicolalde-Morejon et al. 3346 (CIB); 650 m, 24 Sep 2020, L. Martinez-Dominguez et al. 1968 (CIB, MEXU); 500 m, 22 Jan 2001, S. Avendano R. 5375 (MEXU); 518 m, 22 Jan 2001, T. W. Walters 2001 - 39 - D, E (XAL). Mun. San Juquila Vijanos, 1,900 m, 15 Nov 1996, X. Munn et al. 233 (XAL). Mun. San Pedro Sochiapam, 1,682 m, 2 Nov 2016, M. B. Velasco-Pichardo et al. 225 (MEXU). Mun. San Juan Tepeuxila, 1,538 m, 9 May 2008, J. E. Rivera-Hernandez 4289 (MEXU, XAL); 1,538 m, 30 Jul 2008, J. E. Rivera-Hernandez 4380 (MEXU). Mun. Santiago Comaltepec, 1,750 m, 8. Jan 1995, A. Rincon G. et al. 516 (MEXU, MO, XAL); 560 m, 24 Sep 2020, F. Nicolalde-Morejon et al. 3348 - 3355 (CIB); 560 m, 24 Sep 2020, L. Martinez-Dominguez et al. 1973 - 1975, 1980 (CIB, MEXU), 1976, 1978, 1979, 1981 (CIB), 1977 (CIB, MEXU, NY); 1,600 m, 10 Jun 1988, R. Lopez-Luna & G. J. Martin 285 (MEXU), 1,760 m, 26 Jan 1988, R. Torres C. & E. Martinez S. 11345 (MEXU).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
50A0C7C4D7355C5B98D323F9207941D0.taxon	description	Fig. 27 I	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
50A0C7C4D7355C5B98D323F9207941D0.taxon	materials_examined	Type. Mexico. San Luis Potosi: Mun. Rio Verde, 22 Jul 1962, F. Medellin-Leal 1452 ♀ (holotype: SLPM! [acc. # 003530]; isotypes: ENCB! [ENCB 003716], GH! [00003279], MEXU! [MEXU 00162859, MEXU 0053418], MICH! [1192896], US! [00011997]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
50A0C7C4D7355C5B98D323F9207941D0.taxon	description	Description. Stem 10 - 20 cm long, 10 - 15 cm in diameter, semi-hypogeous, erect. Cataphylls 1.8 - 2.5 x 1 - 2 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, partially tomentose at maturity, apex acuminate. Leaves 3 - 27, 95 - 202 cm long, ascending, reddish brown at emergence with whitish gray trichomes, glabrous at maturity. Petiole 11 - 36 cm long, terete, twisted, green in mature leaves, unarmed. Rachis 40 - 77 cm long, terete, twisted, green in mature leaves, unarmed. Leaflets 25 - 46, opposite to subopposite, insertion in one plane, linear, generally longitudinally planar, basally falcate, membranaceous, strongly involute, green with adaxial and abaxial sides glabrous, distal end with entire margins, acute and symmetrical at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 17 - 31.5 x 0.4 - 0.7 cm, 0.8 - 2.3 cm between leaflets; articulations 0.2 - 0.3 cm wide, yellow. Pollen strobili 15 - 19 cm long, 2 - 3.5 cm in diameter, solitary, cylindrical, erect, greenish with reddish brown trichomes at emergence, reddish brown at maturity; peduncle 5 - 8 cm long, 1.5 - 1.8 cm in diameter, tomentose, reddish brown to brown; microsporophylls 0.8 - 1.2 x 0.3 - 0.6 cm, obconic with a non-recurved distal face and a lobate fertile portion, infertile portion 0.25 - 0.35 cm long and rounded with straight horns 0.20 - 0.30 cm long, 0.22 - 0.30 cm and an obtuse angle between the horns. Ovulate strobili 7 - 12 cm long, 5.5 - 7.3 cm in diameter, solitary, cylindrical, erect, green with scarce reddish brown trichomes at emergence, dark green at maturity, acute apex; peduncle 6 - 9 cm long, 0.9 - 1.2 cm in diameter, erect, tomentose, brown; megasporophylls 24 - 49, 5 - 7 orthostichies with 5 - 6 sporophylls per orthostichy, 2.0 - 2.6 x 2.2 - 3.7 cm, with a truncate distal face, horns straight and 0.33 - 0.45 cm long, 1.95 - 2.35 cm between horns with an obtuse angle between the horns. Seeds 2 - 2.8 cm long, 1.8 - 2 cm in diameter, ovate, sarcotesta light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
50A0C7C4D7355C5B98D323F9207941D0.taxon	distribution	Distribution and habitat. Ceratozamia zaragozae is endemic to Mexico in a small mountain range in San Luis Potosi ́ (Fig. 29 C), where it occurs in pine-oak forest on karstic rocks at 1,500 - 1,950 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
50A0C7C4D7355C5B98D323F9207941D0.taxon	etymology	Etymology. The specific epithet refers to General Ignacio Zaragoza, who was a hero in the Battle of Puebla against the French Army in May of 1862.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
50A0C7C4D7355C5B98D323F9207941D0.taxon	discussion	Discussion. Ceratozamia zaragozae and C. norstogii are the only species in the genus with twisted petiole and rachis, but the first has an unarmed petiole, whereas C. norstogii has abundant and robust prickles on the petiole. Additionally, leaflets are membranaceous in C. zaragozae, whereas in C. norstogii they are coriaceous.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
50A0C7C4D7355C5B98D323F9207941D0.taxon	materials_examined	Specimens examined. Mexico. San Luis Potosi: Mun. Rio Verde, 1,700 m, 29 Mar 1984, A. G. Mendoza & L. Vargas 1389 (MEXU); 1,750 m, 20 Sep 1979, A. P. Vovides 435 (XAL); 22 Jul 1962, E. Molseed 34 (MEXU; MICH); 1,860 m, 24 Jan 1994, F. Garcia S. s / n (SLPM); 1,800 m, 22 Jul 1962, F. Medellin-Leal 1451 (SLPM; US); 28 Oct 1965, F. Medellin-Leal s / n (SLPM); 1,900 m, Sep 1994, F. Medellin-Leal s / n (SLPM); 1,956 m, 18 Mar 2016, F. Nicolalde-Morejon et al. 2307 - 2319 (CIB); 1,750 m, 13 Apr 1968, J. Rzedowski 25658 (ENCB; MICH); 1,956 m, 18 Mar 2016, L. Martinez-Dominguez et al. 792, 794 - 796, 798 - 808 (CIB), 793, 797, 799 (CIB, MEXU); 2031 m, 31 Jul 2017, P. Chavez C. et al. 98 (SLPM); 1,532 m, 13 Jan 2001, T. Walters et al. TW- 2001 - 07 (MEXU, XAL). Mun. Zaragoza, 1,869 m, 9 Nov 2012, P. Castillo-Lara et al. 593 (SLPM); 1,847 m, 14 Jul 2016, P. Castillo-Lara et al. 1073 (SLPM).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
4639B9A8F2DA5E839A7B50B9F41ECFCD.taxon	description	Figs 1 D, 23 A-C	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
4639B9A8F2DA5E839A7B50B9F41ECFCD.taxon	materials_examined	Type. Mexico. Chiapas: Northern mountain range, 18 Nov 1998, M. A. Pe ́ rez-Farrera 1732 ♂ (holotype: CHIP [n. v]).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
4639B9A8F2DA5E839A7B50B9F41ECFCD.taxon	description	Description. Stem 12 - 30 cm long, 8 - 15 cm in diameter, epigeous, semi-hypogeous, erect. Cataphylls 2.5 - 6.9 x 1.3 - 4.5 cm wide at the base, persistent, triangular, reddish brown, densely brown tomentose at emergence, glabrous at apex when mature, apex acuminate. Leaves 2 - 17, 46 - 216.5 cm long, descending, light green or brown, glaucous at emergence with whitish gray trichomes, glabrous at maturity. Petiole 23.3 - 111.5 cm long, terete, linear, pink at emergence, yellowish green green at mature leaves; unarmed to armed with 3 - 27 thin prickles 0.08 - 0.15 cm long. Rachis 17.6 - 114.5 cm long, terete, linear, pink at emergence, yellowish green at mature leaves, generally unarmed. Leaflets 6 - 17 pairs, opposite to subopposite, insertion in one plane, oblong to oblanceolate, longitudinally curved abaxially to planar, generally basally falcate, coriaceous, flat, green with adaxial and / or abaxial side glaucous, distal end with entire margins, acuminate (rarely acute) and asymmetrical (rarely symmetrical in apical leaflets) at the apex, attenuate at base, with conspicuous and light-green veins; median leaflets 22 - 37 x 3.9 - 5.6 (7) cm, 4.6 - 11.1 cm between leaflets; articulations 0.5 - 1.5 cm wide, green and yellowish. Pollen strobili solitary (up to 2), 10.8 - 25 cm long, 2.8 - 4.3 cm in diameter, cylindrical, erect, green with blackish trichomes at emergence, yellow-cream with blackish trichomes at maturity; peduncle 5 - 13 cm long, 1.5 - 1.8 cm in diameter, tomentose, light brown; microsporophylls 0.9 - 1.4 x 0.7 - 0.9 cm, discoid with a non-recurved distal face and a deeply lobate fertile portion, infertile portion 0.35 - 0.40 cm long and linear with straight horns 0.30 - 0.45 cm long, 0.73 - 0.80 cm and an acute angle between the horns. Ovulate strobili 15 - 26 cm long, 7.5 - 9.5 cm in diameter, solitary, cylindrical, erect, green with reddish brown trichomes at emergence, green with brown to blackish trichomes at maturity, acute apex; peduncle 5 - 18.5 cm long, 1.6 - 2 cm in diameter, pendulous and erect, tomentose, light brown; megasporophylls 30 - 56, 6 - 8 orthostichies with 5 - 7 sporophylls per orthostichy, 1.5 - 2.0 x 3.5 - 4.0 cm, with a prominent distal face, horns curved to straight and 0.70 - 0.90 cm long, 0.92 - 1.56 cm between horns with a right angle between the horns. Seeds 2 - 2.8 cm long, 1.8 - 2 cm in diameter, ovate, sarcotesta whitish pink at emergence, light brown at maturity.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
4639B9A8F2DA5E839A7B50B9F41ECFCD.taxon	distribution	Distribution and habitat. Ceratozamia zoquorum is endemic to the northern mountains of Chiapas, Mexico (Fig. 29 D), where it occurs on karstic outcrops in evergreen tropical forest and oak forest between 500 and 1,150 m.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
4639B9A8F2DA5E839A7B50B9F41ECFCD.taxon	etymology	Etymology. The specific epithet was established in honor of the Zoque culture (Perez-Farrera et al. 2001).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
4639B9A8F2DA5E839A7B50B9F41ECFCD.taxon	discussion	Discussion. Ceratozamia zoquorum has oblong and coriaceous leaflets and leaves with scarce thin and short prickles. It belongs to a cryptic taxonomic group, and is geographically close to populations of C. becerrae and C. santillanii, the other two species in this group. The three taxa are distinguishable with the nrITS (Martinez-Dominguez et al. 2017 c). Morphologically, C. zoquorum differs from C. santillanii by its peduncle of ovulate strobili more than 3 cm long.	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
4639B9A8F2DA5E839A7B50B9F41ECFCD.taxon	materials_examined	Specimens examined. Mexico. Chiapas: Mun. Solosuchiapa, 530 m, 23 Jan 2014, F. Nicolalde-Morejo ́ n et al. 1931, 1932 (CIB); 550 m, 23 Jan 2014, F. Nicolalde-Morejo ́ n et al. 1933 - 1935 (CIB); 682 m, 24 Jan 2014, F. Nicolalde-Morejo ́ n et al. 1936 - 1947 (CIB); 531 m, 23 Jan 2014, L. Marti ́ nez-Domi ́ nguez et al. 1 - 5 (CIB); 500 m, 23 Jan 2014, L. Marti ́ nez-Domi ́ nguez et al. 6 - 14 (CIB); 682 m, 23 Jan 2014, L. Marti ́ nez-Domi ́ nguez et al. 15 - 34 (CIB); 520 m, 17 Apr 1996, M. A. Pe ́ rez-Farrera 905 (CH, CHIP, HEM); 520 m, 16 Apr 1996, M. A. Pe ́ rez-Farrera s / n (HEM); 19 Jan 2001, S. Avendan ̃ o 5216 (MEXU); 531 m, 20 Jan 2001, T. W. Walters 2001 - 2028 - A (XAL). Mun. Tila, 1,135 m, 16 Jul 2021, F. Nicolalde-Morejon et al. 3698 - 3702 (CIB); 1,135 m, 16 Jul 2021, L. Martinez-Dominguez et al. 2326 - 2330 (CIB).	en	Martinez-Dominguez, Lili, Nicolalde-Morejon, Fernando, Vergara-Silva, Francisco, Stevenson, Dennis Wm. (2022): Monograph of Ceratozamia (Zamiaceae, Cycadales): an endangered genus. PhytoKeys 208: 1-102, DOI: http://dx.doi.org/10.3897/phytokeys.208.80382, URL: http://dx.doi.org/10.3897/phytokeys.208.80382
